Cretaceous Wildfires and Their Impact on the Earth System

Cretaceous Research 36 (2012) 162e190

Contents lists available at SciVerse ScienceDirect

Cretaceous Research

journal homepage: www.elsevier.com/locate/CretRes

Review paper Cretaceous wildﬁres and their impact on the Earth system

Sarah A.E. Brown a, Andrew C. Scott a,*, Ian J. Glasspool b, Margaret E. Collinson a,c a Department of Earth Sciences, Royal Holloway University of London, Egham Hill, Egham, Surrey TW20 0EX, UK b Department of Geology, Field Museum of Natural History, 1400 S. Lake Shore Drive, Chicago, IL 60605, USA c Department of Palaeontology, Natural History Museum, Cromwell Road, London, SW7 5BD, UK a r t i c l e i n f o a b s t r a c t

Article history: A comprehensive compilation of literature on global Cretaceous charcoal occurrences shows that from Received 1 April 2011 the Valanginian on throughout the Cretaceous, terrestrial sedimentary systems frequently preserve Accepted in revised form 7 February 2012 charcoal in abundance. This observation indicates that fires were widespread and frequent and that the Available online 31 March 2012 Cretaceous can be considered a “high-fire” world. This increased fire activity has been linked to elevated atmospheric oxygen concentrations, predicted as in excess of 21% throughout this period and 25% during Keywords: some stages. This extensive wildfire activity would have affected the health, composition, and structure Charcoal of the vegetation and, through habitat loss, probably the fauna. For these reasons, fire activity should be Inertinite Polycyclic aromatic hydrocarbons (PAHs) taken into account in Cretaceous vegetation and climate models. Major changes in vegetation occurred Plant fossils during the Cretaceous. In particular, the angiosperms rose to dominance. Some early angiosperms are Angiosperms interpreted as being of weedy form and as having thrived in disturbed environments. Fires may have Climate promoted angiosperm diversification and spread through their role in environmental perturbation. The Post-fire erosion significant number of charred angiosperm mesofossil assemblages described from the late Early Creta- Oceanic anoxic events (OAEs) ceous supports this hypothesis. Additionally, it can be speculated that severe fires during the Cretaceous would have engendered increased levels of runoff and erosion leading to the mobilization of significant amounts of phosphorous into marine settings. This phosphorous runoff would have contributed to oceanic planktonic blooms and their associated anoxic events. Fire activity remained prevalent into the Late Cretaceous. New data on the distribution of charcoal in the Campanian of Dinosaur Provincial Park, Canada indicate extensive charcoal deposits over a 1.7 myr interval and suggest that some catastrophic bone bed accumulations may have been the result of post-fire erosion-depositional systems. Ó 2012 Elsevier Ltd. All rights reserved.

1. Introduction driven the search for new Cretaceous charcoal localities. Large numbers of studies on Cretaceous angiosperms have been under- Wildfire is an important part of the Earth system today taken over the past 30 years (for a review see: Friis et al., 2006), but (Bowman et al., 2009), but the role of this process in deep time is the key Early Cretaceous floras containing flowers and reproductive just beginning to be explored (Scott, 2000; Berner et al., 2003; structures contain charcoalified mesofossil assemblages (Friis et al., Pausas and Keeley, 2009; Belcher et al., 2010a, b; Glasspool and 2006). Despite these advances, we still have little information on Scott, 2010; Harrison et al., 2010). To date, there have been few the nature of fire-prone vegetation in the Cretaceous, or on how fire attempts to assess the nature and extent of wildfire in the Creta- affects, and is affected by, changes in fire systems through the ceous (e.g., Scott, 2000). Cretaceous. A series of papers in the 1970s and 1980s on the anatomy of The Cretaceous is particularly significant in that it represents charred Cretaceous plants (Alvin, 1974; Scott and Collinson, 1978; a period of high (but falling) CO2 (Haworth et al., 2005) and Harris, 1981) alerted palaeobotanists to this preservation type and a greenhouse climate (Spicer, 2003; Brentnall et al., 2005). The opened the way for more intensive taxonomic investigations. climate has been shown to vary though the Cretaceous with However, it was the discovery of charred flowers (Friis and Skarby, intervals of either significant aridity or rainfall (Spicer, 2003). 1981) and other reproductive structures that have subsequently Spatial differences in climate are also documented. Studies using fossils from coastal regions and continental interiors indicate distinct climatic differences between the two (Spicer et al., 2008). * Corresponding author. Tel.: 44 1784443608. Models agree (Fig. 1) that during the Early Cretaceous atmospheric þ E-mail address: [email protected] (A.C. Scott). oxygen levels rose sharply, subsequently declining gradually

Fig. 1. Modelled (Bergman et al., 2004; Berner, 2009) and calculated (Glasspool and Scott, 2010) atmospheric oxygen concentrations in the Cretaceous with the global change in vegetational composition (after Niklas et al., 1985; Crane and Lidgard, 1989). Time scale after 2010 ICS/IUGS.

(Bergman et al., 2004; Berner, 2009; Glasspool and Scott, 2010). are specific to different plant groups and to fire temperature This large rise may have suppressed the effects of climate on fire (Simoneit, 2002). Some of the gases may recombine and condense activity. The middle to later Cretaceous was also a time of dramatic to form soot (Pyne et al., 1996). The morphology of the soot may be vegetational change (Niklas et al., 1985; Crane and Lidgard, 1989; characteristic of different fuel sources (Harvey et al., 2008). Lastly, Lidgard and Crane, 1990; Lupia et al., 1999) (Fig. 1). In this paper we charcoal is a residual product of pyrolysis (Scott, 2010). Micro- consider the occurrence of charcoal in Cretaceous terrestrial sedi- charcoal, PAHs, soot and water vapour may occur in a fire’s ments and interpret the role of wildfire in the Earth system at this smoke plume (Fig. 2) while all of a fire’s products may be preserved time, reviewing examples from the literature together with new in sediments (Finklestein et al., 2005). However, it is typically larger data. We consider the role of post-fire erosion and other impacts charcoal fragments (mesoscopic, 125 mme1 mm and macroscopic, not only upon the terrestrial system but also on the marine system. >1 mm; Glasspool and Scott, in press: note Scott, 2010 used 180 mm as a lower limit) that are produced predominantly by surface fires, 2. Recognition of wildfire products in the Cretaceous and which are most commonly studied from the fossil record (Scott, 2010). It is meso- and macro-charcoal that provides most evidence Wildfires may generate a range of products that have the of Cretaceous wildfires. In particular, charcoal that preserves potential to be preserved in the fossil record: charcoal, soot and anatomical data allows taxonomic identification and permits an pyrolitic polycyclic aromatic hydrocarbons (PAHs) (Fig. 2). Of these improved understanding of the nature of fire systems in deep time charcoal offers the best means of studying fires in deep time (Scott, (Scott, 2010). Within coals charcoal is petrographically referred to 2010). Burning is an oxidation reaction and is self propagating, as inertinite (Scott and Glasspool, 2007) and this has been used as providing the heat necessary for further pyrolysis. Gaseous a proxy for the level of atmospheric oxygen (Glasspool and Scott, compounds released by burning include PAHs. These compounds 2010). 164 S.A.E. Brown et al. / Cretaceous Research 36 (2012) 162e190

Fig. 2. Fire products from surface and crown fires. Crown fires produce predominantly micro-charcoal together with soot and PAHs (polycyclic aromatic hydrocarbons). More macro-charcoal (together with some micro- and meso-charcoal, soot and PAHs) is often produced by lower temperature surface fires (figure modified from Scott, 2000).

3. Occurrence of charcoal in Cretaceous sediments indicates. The charcoal assemblages from this interval show a range of taxonomic diversity (Tables 2, 3, Figs. 8e10). The early Late Charcoal records are plotted on palaeogeographic maps (Fig. 3) Cretaceous charred mesofossil record is particularly important and and the data from which these were derived is given in Tables 1e3. affords the earliest evidence of several angiosperm clades; Pre-Barremian reports of Cretaceous charcoal are dominated by the however, by the late Late Cretaceous gymnosperm wood is perhaps Wealden facies of Europe (Figs. 4e7). However, there are also the most widely reported charcoalified fossil type (Tables 2, 3) reports of charcoal from the Shubenacadie Basin of Nova Scotia, the though angiosperm reproductive structures are reported in Crimean Peninsula of Ukraine, the Algoa Basin of South Africa and assemblages right up until the latest Maastrichtian (Table 3). Maiya in southwest Madagascar (Table 1 and references therein). Despite longstanding claims for global fires at the KeP boundary The record of inertinite ( fossil charcoal; Scott and Glasspool, (e.g., Wolbach et al., 1990), a number of studies examining the ¼ 2007) in coal from this interval is rather more spartan, with only abundance and distribution of charcoal (Belcher et al., 2003, 2005), the coals of the Valanginian Bickford Formation containing greater the morphology of soot (Harvey et al., 2008) and the nature and than 10% charcoal (Table 1). These earliest Cretaceous fires do not distribution of PAHs (Belcher et al., 2009) indicate that the data do seem to have become abundant until the Valanginian and then not support such a conclusion (Belcher, 2009). occurred throughout the Wealden (Alvin, 1974; Batten, 1974, 1998). New impact models no longer indicate the range of tempera- These fires appear to have burned both fern prairies and large tures required to create widespread and intense fires (Goldin and coniferous trees (Harris, 1981; Batten, 1998; Collinson et al., 2000), Melosh, 2009). Some recent authors continue to refer to KeP fires the charcoal assemblages either being dominated by fern organs, (e.g., Kring, 2007), while others follow the change of view (Schulte a plant group less well represented in later assemblages, or by et al., 2010). conifer wood (Figs. 4e7). From the Barremian until the end of the Early Cretaceous the 4. Implications for the Cretaceous Earth system and biota diversity of plants preserved as charcoal, the range of organs preserved and the geographic distribution across which they are 4.1. The concepts of fire regimes and fire systems reported all increase (Table 1). Despite these increases, fires in the polar biome appear to remain rare during this interval (Falcon-Lang A fire regime is a concept applied to modern ecosystems and et al., 2001) and may have been initiated by volcanic activity. Of takes into account the pattern of fire frequency, severity (see particular note are the Portuguese and the North American Keeley, 2009) and spatial extent (Gill, 1975; Bond and Keeley, 2005) assemblages of this age, which are of prime taxonomic importance (Fig. 11). There is often considerable misunderstanding over the as they preserve many early angiosperm reproductive organs, some terms fire intensity and fire/burn severity. Fire intensity is charred, some lignitic (Friis et al., 2006, 2011, Table 1). These a measure of the energy released, while fire severity measures structures have remained uncrushed because of their low organic matter loss (Keeley, 2009, Fig. 11); neither is easily esti- compaction so preserve fine anatomical details that have allowed mated from the fossil record. In the geological record, where our key insights into the evolution and diversification of the flowering data come predominantly from charcoal, we are limited to plants (Friis et al., 2006, 2011). Also worthy of note is the apparent a consideration of the nature of what has been burnt, and in some frequent association of charcoal with deposits of amber (Brasier cases fire temperature (estimated from charcoal reflectance) (Scott, et al., 2009), fire damage probably having resulted in significant 2010). The influence of fire on the structure of vegetation and resin generation. However, as Brasier et al. (2009) remarked, this dominant plant traits varies depending on the fire regime (Bond association is scarcer in post-Cretaceous deposits. Therefore, this and Scott, 2010). However, fire regimes depend on the synergy association may in part reflect the prevalence of fire during the between external physical factors and the properties of vegetation Cretaceous. (Bond and Scott, 2010). For example, climate may bring about In the Late Cretaceous there is evidence of fire on every conti- changes in plant productivity and flammability that may influence nent (Tables 2, 3, Fig. 3). However, the majority of data comes from the amount and accumulation of fuel. Recent experiments by the Northern Hemisphere, in particular Europe and North America Belcher et al. (2010a) show that the nature of leaf type may influ- with a paucity of data from the vast bulk of the globe. Occasional ence flammability. Whitlock et al. (2010) have introduced the inertinite data from coals exist (e.g., the Coniacian Agwu and concept of the super fire regime. This takes account of the longer Maastrichtian Mamu formations of Nigeria, and the time scale of the geological record, although still considering CampanianeMaastrichtian Pike River Coalfield of New Zealand; thousands rather than millions of years. The super fire regime is Tables 2, 3) and, particularly those from Nigeria, suggest that fires described as “the characteristic nature of fire within a biome, should have been more common than the current mesofossil record integrating all possible variations in climate, and human influences Fig. 3. Geographic distribution of charcoal mesofossil assemblages and inertinite (charcoal in coal) at three Cretaceous time intervals (see Tables 1e3 and Glasspool and Scott, 2010 for data). Plate reconstructions by C.R. Scotese (Paleomap project), with permission. A, BerriasianeAlbian data, plotted on a 120 Ma map. B, CenomanianeSantonian data, plotted on a 100 Ma map. C, CampanianeMaastrichtian data, plotted on a 80 Ma map. 1

Table 1 6 6 Localities of Early Cretaceous (Berriasian eAlbian) sediments yielding charcoal, including assemblag es with charred angiosperm fertile organs and records of quantitati ve inertinit e data from coals: see Fig. 3.

Early Cretaceous (Berriasian eAlbian) Country Locality Stratigraphy Age Lithology and Charred vegetation Uncharred vegetation Comments References environment of deposition England Portland, Dorset Lower Purbeck Jurassic-Cretaceous Palaeosol- black Charcoal present, affinity Silicified conifer Silicified in situ tree Francis, 1983; Fm, Great Dirt boundary carbonaceous marl unspeci fied. wood & shoots. stumps occur in the Watson and Alvin, 1996 Bed with limestone Lower Dirt Bed along pebbles deposited with conifer branches. at the edge of There is no a hypersaline lagoon. reported charcoal from this bed. Germany Wealden Basin Wealden Berriasian Coal. Inertinite (mmf) 6.9%, Average of 11 samples range: <10% reported in Glasspool and Scott (2010) England Hastings Ashdown Fm latest Berriasian- Amber in Only conifer This amber deposit Brasier et al., 2009 early Valangin ian “charcoal (fusain) wood. may owe much to lignites ”. fire as resins “flowed S.A through the carbonized .E. tracheids and into broader Brown fissures” following burning. Charcoal “is also known to be associated with other et early amber nodules e.g. al. /

the Hauterivian of Lebanon; Cretaceous Barremian of the Isle of Wight; Barremian to Maastrichtian of Spain) although it tends to

be scarcer in much younger Resear amber deposits ”.

Ukraine Crimean lower Valanginian Shallow marine. Not speciﬁed. Grocke et al., 2005 ch

Peninsula, 36

Kacha River, (20

Rezanaya 1 2) Mountain 1 Australia Surat Basin Orallo Fm Valanginian Coal. Inertinite (mmf) 1.0%, range: Data from Glasspool 62 e

<10% and Scott (2010) 1 9

Canada British Columbia Bickford Fm Valanginian Coal. Inertinite (mmf) 18.3%, range: Data from Glasspool 0 10e20% and Scott (2010) Canada British Columbia Minnes Gp, Valanginian Coal. Inertinite (mmf) 1.0%, range: Data from Glasspool Gorman Creek <10% and Scott (2010) Fm South Africa Algoa Basin, Kirkwood Fm middle-upper Mudstone Wood (up to 15 mm long). Impressions of bryophytes, Gomez et al., 2002 Sundays River, Valanginian palaeosoils and ferns, cycads, bennettites, Mfuleni farm coarse conifers & unassigned fluvial sandstones. gymnosperms. Also conifer & cycadalean leaves in amber. Ukraine Crimean upper Valangin ian Shallow marine. Not specified, but Grocke et al., 2005 Peninsula, undifferentiated tracheids Kacha River, illustrated from this interval. Rezanaya Mountain Canada Nova Scotia, Chaswood Fm Valanginian- Sands & clays. Ferns & conifer wood. Conifer wood, leaves Evidence of post-fire soil Scott and Stea, 2002; Shaw Pit, Hauterivian & cones & lycopsid erosion (quartz grains in Falcon-Lang et al., 2007 Shubenacadie megaspores. lignite horizons). Basin Canada Nova Scotia, Chaswood Fm Valanginian- Coarse grained, Gymnosperm wood Ferns & bennettite cuticle. Falcon-La ng et al., 2007 Bailey Quarry, Hauterivian pebbly (w1 cm3) -Taxodioxylon Shubenacadie sandstones & clays. (predominates), Basin Cupressinoxylon (present). Ukraine Crimean lower Hauterivian Shallow marine. Not specified. Grocke et al., 2005 Peninsula, southern slope of Belaya Mountain Madagascar SW Madagascar, Hauterivian Ferns (Gleichenimorpha Bennettites (dominant), Despite a diverse flora Appert, 2010 Manja monostigma, Phlebopteris ferns, spehnopsids, apparently dominated by dunkeri, hepatophytes, benettites, but also including Weichselia reticulata ). pteridosperms, conifers, angiosperms, the only charred & angiosperm leaves (few). remains reported are those of ferns. England Isle of Wight, Wessex Fm Hauterivian- Mudstones Charred wood (<1 cm3) - Silicified Araucarioid Abundant charred mesofossils, Collinson et al., 2000 Hanover Point Barremian deposited Podocarpaceae wood, leafy conifer but uncharred araucarioid Assemblage along river (predominates), shoots & termite coprolites. wood is three times as channel margin. Pseudofrenelopsis (rare). common as charred wood. Assemblage derived from an environment dominated by S.A low diversity conifers, some .E. of which were large trees. Brown England Isle of Wight, Sheperds Chine Hauterivian- Low energy Gymnosperm wood Large uncharred Abundant charred mesofossils. Alvin, 1974; Collinso n Sheperds Chine Mbr, Vectis Fm Barremian shallow coastal (<0.5 mm3 to >1 mm3) e component including The siltstones occur in runnels et al., 2000 (See:

Assemblage lagoon Aff. Pinaceae, Taxodiaceae wood, plus rarer (Fig. 5D, E), possibly as a Fig. 5 this paper) et mudstones, or Cupressaceae. Ferns megaspor es, bennettite result of rapid deposition al. ﬁ / siltstones (predominate) -Weichselia & cycad cuticles, & following post- re erosion. Cretaceous & sandstones (Fig. 5C) & other taxa (Fig. 5B). arthropod cuticle & (Fig. 5A,B). ﬁsh vertebrae. England Surrey, Clock Lower Weald Hauterivian Mudstone. Wood, fungal hyphae, A highly diverse The charcoal assemblage Batten, 1998

Pit south of Capel Clay Fm fern fronds (including assemblage of seeds occurs in facies with dinosaur Resear Weichselia ), gymnosperm & gymnosperm organs. teeth & bones. The assemblage

seeds, conifer shoots may reﬂect post-ﬁre erosion. ch

& arthropod remains. 36

England Surrey, Upper Weald Barremian Mudstone. Wood, fungal hyphae, A highly diverse The charcoal assemblage Batten, 1998 (20

Smokejacks Clay Fm fern fronds (including assemblage of seeds occurs in facies with dinosaur 1 Brickworks, Weichselia ), gymnosperm & gymnosperm organs. teeth & bones. The 2) 1 Ockly seeds, conifer shoots assemblage may reﬂect post- 62

ﬁ e

& arthropod remains. re erosion. 1 9

England Isle of Wight, Wealden Marls, earliest Barremian Amber in Unidenti fied wood (fusain). Conifers (abundant) Nicholas et al., 1993 0 Chilton Chine Wessex Fm lignite formed probably Brachyphyllum , as a channel- and 21 miospore species. lag deposit. England Surrey, Beare Upper Weald Barremian Broad river Only ferns: Weichselia None reported. Harris, 1981 Green Brick Pit, Clay Fm channel facies. (abundant), Gleichenites Holmwood & Phlebopteris dunkeri (less common). Belgium Mons Basin, Hautrage Fm Barremian Floodplain Wood, coniferous twigs Ginkgoalean fruit cuticle, Plant composition varies Gomez et al., 2008, Danube- clays, silts & seeds, unidenti fied conifer twigs, leaves, within each of the seven beds in press Bouchon Quarry & sands. reproductive structures, seeds, cones & scales reported at this locality. Some ferns (Fig. 7) & fern litter & scales & cones of of the channel sands contain (Fig. 6). unknown affinity. rip-up clasts of charred litter dominated by ferns (Fig. 6). (continued on next page) 1 6 7 1

Table 1 (continued ) 6 8

Early Cretaceous (Berriasian eAlbian) Country Locality Stratigraphy Age Lithology and Charred vegetation Uncharred vegetation Comments References environment of deposition U.S.A Virginia, Drewry ’s Potomac Gp Barremian e Aptian Clay balls Not specified. Not specified. Charcoal is present at this Pedersen et al., 1993; Bluff, southeast embedded in locality along with lignitised Crane and of Richmond a sand and compressions. It is unclear Herendeen, 1996 gravel bed. which flora has been charred. Leaf bed is Flora is contained within clay contained balls and leaf beds. within a thin The clay balls contain fern clay bed higher fragments, gymnospermous in the section leaves, twigs & seeds. than the clay The leaf bed contains balls. angiosperm leaves, fern fronds, conifer shoots & cycad leaves. U.S.A Virginia, Dutch Patuxent Barremian eAptian Fluviatile/ Not specified. Not specified. Charcoal is present at this Pedersen et al., 1993; Gap, southeast Formation, deltaic cross locality along with lignitised Crane and S.A of Richmond Potomac Gp bedded sands, compressions. It is unclear Herendeen, 1996 .E. laminated clays which flora has been charred. Brown and silt, gravel Vegetation includes 12 beds. angiosperm leaf types (magnoliid affinity). Conifers, et bennettitalean leaves, al. /

ginkgoalean leaves and fern Cretaceous fragments are all present. Portugal Torres Vedras, Base of late Barremian-early Sandy lignite. Angiosperm ﬂowers, Megaspores & pollen Charred angiosperm ﬂora less Friis et al., 2004 Lusitanian Basin Almargen Fm Aptian fruits & seeds. within coprolites diverse than other early

Unidenti fied fragments. (small component). Cretaceous Portuguese localities. Resear Portugal Catefica, late Barremiane Fluviatile cross- Angiosperm flowers, Rare lignitic compressions. Friis et al., 1999

western margin Aptian bedded sands fruits & seeds, ferns, ch

of Runa Basin with organic liverworts & conifer twigs. 36

horizons (20

& clay beds. 1 2) Canada Alberta & Gething Fm Aptian Coal. Inertinite (mmf) 19.1%, Average of 68 samples 1 British Columbia range: 10e20% reported in 62 e

Glasspool 1 9

and Scott (2010) 0 Portugal Famalicão, late Aptian Organic rich clay. Angiosperm fruits, Lignitised angiosperms. Most diverse floral assemblage Friis et al., 1994, western seeds & flowers, from the early Cretaceous 1999, 2000; Portuguese Basin Cheirolepidiaceous of Portugal. Eriksson et al., 2000 conifers (rare). Portugal Vila Verde, ENE “Arenitos de late Aptian-early Alternating Angiosperm flowers, Impressions of conifer Friis et al., 2006, 2010; of Figueria de Carrascal ” Albian clay & silt layers. leaves & seeds, twigs, ferns Friis and Pedersen, 1996 Foz area complex unspeci fied wood. & angiosperm leaves (rare). Portugal Vale de Agua, Famalicão Mbr late Aptian-early Clay. Angiosperm fruits Lignitised flowers von Balthazar Lusitanian Basin of Figueria de Albian & seeds, Magnoliid & pollen organs. et al., 2005; Friis Foz Fm & Ranunculalean flowers. et al., 1997, 1999, Charred conifer cones 2006; Pedersen and twigs. et al., 2007; Mendes et al., 2010 Portugal Buarcos, Beira “Arenitos de late Aptian-early Cross bedded Angiosperm flowers None reported. Gymnosperm twigs of Friis et al., 1997, Litoral region Carrascal ” Albian coarse sandstone & fruits (Chloranthaceae). Cheirolepidiaceae affinity - 1999, 2000 complex with fluviatile not clear if these are charred. or lacustrine intercalated layers of clay & silt. U.S.A. Texas, Hood County Twin Mountains Aptian-Albian Fluvially Wood (Cheirolepidiaceae). Silicified logs Extends geographical extent Axsmith and Jacobs, Fm derived well (Cheirolepidiaceae), of Frenelopsis ramosissima 2005 cemented gymnospermous by 2100 km within the U.S.A.. sandstones. shoots & cones. Dinosaurs found within this formation. U.S.A Arkansa s, north Holly Creek Fm Aptian-Albian Clay bed. Charcoali fied wood is Uncharred wood and Axsmith, 2006 of Nashville present, affinity has not twigs, silicified wood. been specified. Brazil Araripe Basin, Romualdo Mbr, Early Albian Lacustrine Woody gymnospermous Calcitized plant debris Charcoal is contained within Martill et al., 2012 Caatinga Santana Fm silty shales fragments. including cones from the carbonate concretions. with carbonate cycadaceans and 3D fish and rare pterosaur s concretions. Equistetales stems. have also been observed Rare calcitized logs within the concretions. are present. England Bedfordshire, 3m Below the early Albian Tidal flat, Gymnosperm wood None reported . Herendeen and Skog, Munday ’s Hill Gault Fm organic-rich (abundant), fern pinnules 1998 Quarry near clays & silts & rachis (Gleichenia). Leighton Buzzard from near an estuary mouth. Spain Cantabria Las Penosas Fm early Albian Amber within Charcoali fied wood is Alvinia cones and A few amber pieces contain Najarro et al., 2010 S.A carbonaceous present, however affinity Frenelopsis branching charcoa lified plant fibres. .E. shales, siltstones, has not been specified. shoots. Charred Weichselia occurs Brown and wavy/ in adjacent beds to those lenticular containing amber.

sandstone et layers. Interpreted al. /

to have been Cretaceous deposited in an estuarine bay system with

small bayhead Resear deltas.

Antarctica Kerguelen Lith. Unit IV early Albian Clayey siltstone Only wood (Podocarpoxylon ). Fires may have been initiated Francis and Cofﬁn, 1992 ch

Plateau, Site with volcaniclastics by volcanism. 36

750, Raggatt Basin and coal. (20 ﬂ

U.S.A. Virginia, Potomac Gp early-middle Albian Clay & silt. Angiosperm fruits, wood Compressed lignitic Most diverse mesofossil ora Herendeen, 1991a, 1 Puddledock & ﬂowers (Aff. ﬂowers & fruits. from the Early Cretaceous of b; Srinivasan, 1992; 2) 1 in Prince County Magnolialean or Lauralean). North America. Crane and Herendeen, 62 e

1996, Friis et al., 1994; 1 9

von Balthazar et al., 2007 0 U.S.A. Virginia, Bank Potomac Gp early-middle Albian Sandstones Not specified. Not specified. Charcoal is present at this Crane et al., 1993; near Brooke with intercalated locality along with lignitised Crane and claystones and compressions. It is unclear Herendeen, 1996 siltstones. which flora has been charred. Angiosperm leaves, conifer shoots and leaves and fragmentary ferns are present. U.S.A. Virginia, Quantico Potomac Gp early-middle Albian Not specified. Not specified. Charcoal is present at this Crane and locality along with lignitised Herendeen, 1996 compressions. It is unclear which flora has been charred. Vegetation includes gymnospermous leaves and shoots, conifers, cycadophyte, pteridophyte; along with angiosperm leaves. (continued on next page) 1 6 9 1 Table 1 (continued ) 7 0

Early Cretaceous (Berriasian eAlbian) Country Locality Stratigraphy Age Lithology and Charred vegetation Uncharred vegetation Comments References environment of deposition Spain Valle del Rio Martin Esucha Fm, La Albian Floodplain Unspeci fied elements of Impressions & Sender et al., 2005 Orden Mbr near channel the compression and compressions of levee; clay-rich, impression flora. Brachyphyllum with fine-grained cones (abundant), ferns sandstone. (Cladophlebis & Weichselia reticulata ), Zamites & Cycadales, Australia Burgowan Coal Albian Coal. Inertinite (mmf) 9.0%, range: Data from Glasspool <10% and Scott (2010) Canada Alberta & Gates Fm Albian Coal. Inertinite (mmf) 29.9%, range: Average of 19 samples British Columbia >20% reported in Glasspool and Scott (2010) Canada British Boulder Albian Coal. Inertinite (mmf) 45.9%, range: Data from Glasspool Columbia, Peace Creek Fm >20% and Scott (2010) S.A River .E. Canada Mannvile Gp Albian Coal. Inertinite (mmf) 23.4%, range: Average of 19 samples Brown >20% reported in Glasspool and Scott (2010) Spain Albian Coal. Inertinite (mmf) 22.3%, range: Average of 14 samples et >20% reported in Glasspool al. /

and Scott (2010) Cretaceous U.S.A. Atlantic Sample 8030 Albian Coal. Inertinite (mmf) 17.8%, range: Data from Glasspool Continental 10e20% and Scott (2010) Slope, New Jersey

Antarctica Alexander Island Triton late Albian Water-lain Conifer wood (dominant) Fires considered rare in the Falcon-Lang et al., 2001 Resear Point Fm tuffs within Araucariopitysm & polar biome and may have

meander ebelt Podocarpoxylon , podocarp been initiated by volcanism. ch

association. twigs, scale leaves (cf. 36

Brachyphyllum, (20

Pagiophyllum ) and 1 2) seed-coats. 1 U.S.A. Maryland, West Patapsco Fm latest Albian Abandoned Angiosperm ﬂowers. Lignitic and First evidence of the buxaceous Crane and Upchurch, 62 ﬂ e Brothers, channel silts, compressed owers. lineage. Angiosperm 1987; Friis et al., 1988; 1 9

Prince Georges clays & cross- & gymnosperm wood, conifer Drinnan et al., 1991 0 County bedded sands. cones, shoots & seeds are also present. It is unclear whether these are charred or lignitised. Table 2 Localities of early Late Cretaceous (Cenomanian eSantonian) sediments yielding charcoal, including assemblag es with charred angiosperm fertile organs and records of quantitati ve inertinit e data from coals: see Fig. 3.

Early late Cretaceous (Cenomanian eSantonian) Country Locality Stratigraphy Age Lithology Charred vegetation Uncharred vegetation Comments References U.S.A. Kansas, Ellsworth Dakota Fm Albiane Fine sands, Undifferentiated wood Charcoal accounts for between w1e11% Wang, 2004 and Cloud counties Cenomanian shales and clays. (dominant). by weight of the sediments studied. 4899 mesofossils (267 morphotypes) including angiosperm flowers, fruits & seeds, conifers & pteridophytes. Fossils lignitic or charred. Assemblages dominated by gymnosperms, but angiosperms were common & diverse. U.S.A. Alaska, Kukpowruk Kukpowruk Albiane Siltstones and Charcoali fied wood Ironstone preserved twigs. Spicer and Herman, 2001 River Fm Cenomanian nudstones fragments. interpreted as representing woody mires. Australia Queensland, Winton, late Albiane Unspeci fied, probably Bennettitales, Cycadales and Pole and Douglas, 1999 Eromanga Basin Mackunda, Cenomanian wood in associat ion ginkgophytes. S.A & Allaru Fms with Ptilophyllum . .E. Argentina Bajo Comision, Kachaike Fm late Albiane Near-shore Wood (unidenti fied). Angiosperm leaves, fern Rare fires. Passalia, 2007 Brown Santa Cruz Provinc e Cenomanian marine facies fronds & branches grading to with Brachyp hyllum deltaic and et fluvial facies. al. /

U.S.A. South Central Dakota Fm Late Albiane Grey clays. Fern sporangia & synangia None reported. Provides evidence for Marattiaceaen Hu et al., 2006 Cretaceous Minnesota Cenomanian (Aff. Marattiace ae). ferns post-Jurassic within North America. U.S.A. North Maryland, Latest Albiane Charcoal present, type Not speciﬁed. Platanoid leaves, wood and reproductive Crane and Herendeen, 1996 Bull Mtn Cenomanian not speciﬁed. structures are present at this locality

however it is not speciﬁed if these Resear are charred.

Czech Pecínov Quarry, Peruc- Mid-late Braided river Angiosperm ﬂowers Angiosperm & cycad Varied fossil assemblages within the Eklund and Kvacek, 1998; ch

Republic Bohemian Korycany Fm Cenomanian to estuary & wood, conifer wood leaves, conifer cones, six environmental settings represented Falcon-Lang et al., 2001, 36

Cretaceous Basin, mouth sediments. & twigs, ginkgo foliage & twigs, in this quarry. 2004; Kvacek and Eklund, (20

NW Prague branches, ferns & lycopsids. lycopsids, ferns 2003; Ulicný et al., 1997 1 2) & bennettites (rare). 1 Czech Brník, Bohemian Peruc- Cenomanian Fluviatile/ Abundant angiosperms. Impressions Kvacek and Friis, 2010 62 e

Republic Cretaceous Basin, Korycany Fm Meandering & compressions - 1 9

east of Prague river, sandy fossil type unknown. 0 mudstone. Czech Hloubetín- Hute, Peruc Mbr, Cenomanian Charcoal present, type Compressions and Angiosperms are present, however it is Eklund and Kvacek, 1998; Republic Prague Peruc- not specified. lignitized flora, not clear whether these are charred. Kvacek and Eklund, 2003 Korycany Fm type not specified. Czech Moravia Peruc Mbr Cenomanian Angiosperm fruits and Not specified. Knobloch and Mai, 1991 Republic seeds, including those with Platanaceae affinities. U.S.A. Maryland, Maudlin Potomac Gp, Early Clays. Angiosperm wood & Lignitized flowers Drinnan et al., 1990; Mountain, Elk Neck Elk Neck Beds Cenomanian flowers (Aff. Lauraceae). (Aff. Lauraceae). Herendeen 1991a, 1991b Peninsular USA Colorado, San Juan Dakota Coal Cenomanian Coal. Inertinite (mmf) 42.8%, >20% Data from Glasspool and River Coalfield Scott (2010) USA Utah, Southwestern King Cannel Cenomanian- Coal. Inertinite (mmf) 8.2%, <10% Data from Glasspool and Utah Coalfield Coal Turonian Scott (2010) boundary (continued on next page) 1 7 1 1 Table 2 (continued ) 7 2

Early late Cretaceous (Cenomanian eSantonian) Country Locality Stratigraphy Age Lithology Charred vegetation Uncharred vegetation Comments References Kazakhstan Rudnyy, Kustanay Shet-Irgiz Suite Cenomanian- Lacustrine/ Rare charcoali fications- 3D lignitized angiosperm Frumin and Friis, 1996 Region Novokozyrevsk early Turonian alluvial type not specified. flowers, fruits, seeds, suite sands and clays. cone scales, cones, Lagoonal needles, twigs & kaolinitic fern sporangia. clays & silts. Canada British Columbia, Kaskapau Fm late Coal. Inertinite (mmf) 1.0%, <10% Data from Glasspool and Bullmoose Creek Cenomanian Scott (2010) to middle Turonian New Pitt Island Tupuangi Fm Turonian Mudstone. Conifer leaves, wood None reported . Abundant charred conifers indicate Pole and Philippe, 2010 Zealand (Taxodioxylon ) fire was important in this ecosystem. & ginkgoleans (rare). U.S.A. New Jersey- Sayreville Raritan Fm Turonian Levee/back Angiosperm, fruits, seeds None reported . Oldest known record of Capparales. Crepet and Nixon, 1992; levee/swamp & flowers (Aff. Lauraceae, Fossil Triuridaceae flowers represent Gandolfo et al., 1998, 2002; fluvial silts Triuidaceae, the ‘oldest unequivocal fossil monocots ’. Herendeen et al., 1993; S.A & clays. Chloranthaceae, Nixon and Crepet, 1993 .E. Capparales, & platano ids), Brown (predominate) & ferns (present). U.S.A. New York, Staten Raritan Fm Turonian Lignite in sands. Lignite/wood. Conifer cones, cone In situ combustion of the lignite. Hollick, 1906; et Island, Charleston scales, leafy twigs Hollick and Jeffrey, 1909 al. /

(Kreishcherville) & wood, & Czekanowskia Cretaceous U.S.A. Maryland, Cape Sable, Raritan Fm Turonian Lignite in clay. Lignite/wood. Charcoal in association with amber. Troost, 1821 U.S.A. New Jersey, Crossman South Amboy middle-late Clay. Charcoal is present, Not specified. Angiosperm flowers & fruits with Crane and Herendeen, 1996 Fire Clay Mbr, Turonian type not specified. magnoliid & lauraceae affinity are

Raritan Fm present; however it is not clear Resear whether these are charred.

Czech Slezské Pavlovice Upper Low abundance Not speciﬁed. Knobloch and Mai, 1991 ch

Republic Borehole, Silesia Turonian- angiosperm fruits and 36

Lower seeds. (20

Coniacian 1 2) U.S.A. Utah, Emery Coalﬁeld Mancos Shale, Turonian to Coal. Inertinite (mmf) 16.5%, 10e20% Data from Glasspool 1 Ferron Coniacian and Scott (2010) 62 e

Sandstone Mbr 1 9

Nigeria Middle Benue Trough Agwu Fm Coniacian Coal. Inertinite (mmf) 27.5%, >20% Data from Glasspool 0 and Scott (2010) Czech Klikov Klikov Fm Turonian- Fluvial Angiosperm fruits Leaf impressions Potential charred insect eggs, however Váchová and Kvacek, 2009; Republic Santonian & lacustrin e & seeds belonging to & compressions these may be seeds. Hermanová and Kvacek, sandstones 92 genera including representing 23 species. 2010 & mudstones. Aff. Liriodendron , Saurauia & Sabia. U.S.A. New San Juan River Green Coal Turonian Coal. Inertinite (mmf) 11.0%, 10e20% Data from Glasspool and Coalfield to Santonian Scott (2010) U.S.A. New San Juan River Mesaverde Turonian to Coal. Inertinite (mmf) 14.5%, 10e20% Data from Glasspool and Coalfield Gp, Crevasse Santonian Scott (2010) Canyon Fm Canada Vancouver Island Comox Fm Turonian to Coal. Inertinite (mmf) 18.8%, 10e20% Average of 2 samples Santonian reported in Glasspool and Scott (2010) U.S.A. Utah, Kaiparowits Christensen Coniacian Coal. Inertinite (mmf) 13.0%, 10e20% Data from Glasspool and Plateau Coalfield Coal Zone to Santonian Scott (2010) Japan North-eastern Futaba Gp, early Alluvial fan, Angiosperm flowers Lycopsid megaspores Cornalean fruits indicate ‘minimum Takahashi et al., 1999a, Honshu, Kamikitaba Asamiga wa Santonia n poorly sorted, (Archae fagaceae & fern leaves. age for the early divergence of the 1999b, 2008; Friis et al., Mbr, Ashizawa sandy siltstone. futabensis ), fruits asteroid clade’. 2010; Fm (Cornalean), seeds, leaf fragments & wood. Conifer shoots, pollen cones, cone scales, seeds & fern rachides. U.S.A. Georgia, Upatoi Creek Eutaw Fm Santonia n Charcoal present, type Not specified Crane and Herendeen, 1996 not specified. Antarctica Eastern side of Table late Santonian Marine or Angiosperm flowers, Megaspores. Eklund, 2003; Eklund et al., Antarctic Peninsular Nunatak Fm extremely fruits, seeds & leaves 2004 distal deltaic (Aff. magnoliid, eudicot), distributary/ ferns, conifer shoots, estuarine leaves, pollen cones, mouth very wood & seeds (abundant). fine sandstones & siltstones. U.S.A. Georgia, Allon Gaillard Fm, late Santonain Lower floodplain Angiosperm flowers, Lignitised angiosperm Similar organ assemblage composition Herendeen et al., 1999; Sims Buffalo pond, fruits, leaves & seeds, flowers & leaves, to that observed in a modern heathland et al., 1998, 1999; Scott S.A Creek Mbr microlaminated conifer leaves, cones, conifers, ferns & lycopsids. fire observed in Surrey. Assemblage et al., 2000; Scott, 2010; .E. carbonaceous leafy shoots & mosses. likely to represent a surface fire. Lupia, 2011 Brown clay lens. Charred Gleicheniaceae ferns.

Sweden Åsen, Kristianstad late Santonian Lacustrine Angiosperm flowers Lignitic compressions Evidence of adaptation to insect Crane et al., 1989; Friis 1983, et Basin, north-east finely laminated (Actinocalyx bohrii), of flowers, conifer pollination prior to end Cretaceous 1984, 1985; al. /

Scania, Southern clays, silts & fruits and seeds. twigs, cone & ferns. (Actinocalyx bohrii). Endress and Friis, 1991; Cretaceous Sweden sands. Taxodiaceaous conifers This locality also contains early Herendeen, 1991b; Leng and charred ferns. Campanian charcoa l (see Table 3). et al., 2005; Friis et al., 2011 U.S.A. North Carolina, Black Creek latest Fluviatile sands, Charcoal present, type Lignitized ﬂora, type Assemblage is dominated by lignitized Crane and Herendeen, 1996;

Neuse River cut-off, Formtion Santonia n- silts and clays. not specified. not specified. compressions with some Friis et al., 1988; Frumin Resear Goldsboro. early charcoali fications. Which flora are and Friis, 1996

Campan ian charcoali ﬁed has not been speciﬁed. ch

Angiosperm reproductive organs, 36

leaves and wood are present along (20

with conifer twigs. 1 Hungary Bakony Mtns Ajka Fm Upper Charred angiosperm Not speciﬁed. Knobloch and Mai, 1991 2) 1 Santonia n- ruits & seeds with 62 e

Lower abundant specimens 1 9

Campan ian with Magnoliaceae 0 affinities. Germany Aachen Aachen Fm Upper Charred angiosperm Not specified. Knobloch and Mai, 1991 Santonia n- fruits & seeds. Highly Lower diverse assemblage Campan ian with a high abundance (over 100 specimens). Menispermaceae, Clethraceae & Cyrillaceae dominate. Holland South-Limburg Aachen Fm Upper Charred angiosperm Not specified. Knobloch and Mai, 1991 Santonia n- fruits & seeds. Highly Lower diverse assemblage Campan ian with a high abundance (over 100 specimens). Menispermaceae, Clethraceae & Cyrillaceae dominate. 1 7 3 1 7

Table 3 4 Localities of late Late Cretaceous (Campanian eMaastrictian) sediments yielding charcoal, including assemblages with charred angiosperm fertile organs and records of quantitati ve inertinit e data from coals: see Fig. 3.

Late Late Cretaceous (Campa nianeMaastrictian) Country Locality Stratigraphy Age Lithology Charred vegetation Uncharred Comments References vegetation Sweden Åsen, early Campanian Fluviatile cross Angiosperm flowers (Aff. Rare compressions. Charred angiosperms Endress and Friis, 1991; Kristianstad bedded & Saxifragales & Chloranthaceae), dominate with 100 Friis, 1983, 1984, 1985; Basin, north-east laminated sands, fruits & seeds. different taxa & 20 Friis and Pedersen, 1990; Scania, Southern silts & clays. Angiosperm wood - flower types identified. Friis et al., 1986, 2011; Sweden unclear which This locality also Friis and Skarby, 1981., horizon this contains late Santonian 1982; Herendeen, 1991b; belongs to. charcoal (see Table 2). Leng et al., 2005; Schönenberger et al., 2001 U.S.A. SE Arizona Fort Crtittenden Santonian/Campanian Aluvial fan & Charcoal present, type not Not specified. PAH data from this Finklestein et al., 2005 Fm braided river specified. locality indicates shales. variations in intensities of the wildfire. Reflectance data from the charcoal indicates combustion S.A temperatures between .E. 470 and 550 C. Brown U.S.A. Wyoming, Hams Adaville Fm Santonian to early Coal. Inertinite (mmf) 4.9%, Average of 7 samples Fork Campanian <10% reported in Glasspool and Scott (2010) et Portugal Esgueira, north- “Arenitos e Conacian- Maastric tian Alternating Gymnosperm & Abunda nt lignitized Unhcarred component Friis et al., 1992, 2003; al. /

east Aveiro, Beira argilas de Aveiro” sands, silts angiosperm wood. compressions of flowers. dominates the assemblage. Herendeen 1991b Cretaceous Litoral region & clays. Leaf impressions & conifer leaf mats. Portugal Mira, south of “Agilas de Vagos” Campanian e Clay & silt. Angiosperm flowers (Aff. Lignitized flower Angiosperm flowering Friis et al., 1992, 2003;

Aviero, Beira & “Conglomerado Maastrichtian. Myrtales & Fagales), fruits & leaf fragmen ts. structures give evidence Schönenberger et al., 2001 Resear Litoral region de Mira” (abundant), wood of wind pollination.

& gymnosperm wood. ch

U.S.A. Massachusetts, Magothy Fm Campanian Lignitic clay. Diverse assemblage of 3D compactions of Tiffney, 1977, per comm. 36

Matha’s Vineyard gymnosperms & charred gymnospermous (20

fruits & seeds. wood, cones 1 2) & leaves, angiosperm 1 ﬂowers, fruits & seeds. 62 ﬂ e Canada Alberta, Dinosaur Oldman Fm Campanian Palaeochannel Gymnospermous Gymnospe rm fragments. Wide area in uenced This paper 1 9

Provincial Park find sandstones, wood only (125 mm-3cm). by fire & associated 0 mudstones post-fire erosion. & shales. Canada Alberta, Dinosaur Dinosaur Park Fm Campanian Medium Gymnospermous wood Gymnospe rm fragments, Wide area influenced This paper Provincial Park sandstones, only (125 mme3 cm). cones & an unidenti fied by fire. Associated mudstones leaf impression. post-fire erosion & shales. hypothesised to have Interpreted as been involved in the representing accumulation of point bars & dinosaur bone beds. thick overbank facies of meandering rivers. U.S.A. Utah, Sego Coalfield Nelson Fm Campanian Coal. Inertinite (mmf) Data from Glasspool 18.7%, 10e20% and Scott (2010) U.S.A. New Mexico, San Menefee Fm Campanian Coal. Inertinite (mmf) Data from Glasspool Juan Basin 14.0%, 10e20% and Scott (2010) Austria Gosau Fm Campanian Coal. Inertinite (mmf) Average of 24 samples 6.1%, <10% reported in Glasspool and Scott (2010) U.S.A. Wyoming, Green Rock Springs Fm Campanian Coal. Inertinite (mmf) Data from Glasspool River Coalfield 6.3%, <10% and Scott (2010) U.S.A. New San Juan Basin Fruitland Fm Campanian Coal. Inertinite (mmf) Data from Glasspool 16.1%, 10e20% and Scott (2010) U.S.A. Utah, Uinta Coalfield Rock Canyon Coal Campanian Coal. Inertinite (mmf) Data from Glasspool 22.4%, >20% and Scott (2010) U.S.A. Utah Blackhawk Fm Campanian Coal. Inertinite (mmf) Average of 76 samples 12.2%, 10e20% reported in Glasspool and Scott (2010) Canada NW Ellesmere Hasen Point Campanian to Coastal plain/ Only conifer wood (2 cm). None present. Uncharred Falcon-Lang et al., 2004 Island, unnamed Volcanic Unit Maastrichtian peat mire horizon. angiosperm foliage, peninsular ginkgoleans, conifer between Emma shoots & wood Fiord & Audhild Bay (silicified) occur two horizons above. Silicified wood exhibits evidence of disturbance. New Zealand Pike River Coalfield Member s 3 & 4 Campanian to Coal. Inertinite (mmf) Average of 2 samples Maastrichtian 7.9%, <10% reported in Glasspool S.A and Scott (2010) .E. New Zealand Pike River Coalfield Rewanui Coal Campanian to Coal. Inertinite (mmf) Average of 48 samples Brown Measures Maastrichtian 7.2%, <10% reported in Glasspool and Scott (2010)

Austria Vienna Flysch Sievering Fm Campanian e Low diversity and abundance Not specified. Knobloch and Mai, 1991 et Maastrichtian of angiosperm fruits & seeds. al. fi / Czech Rupublic Flysch, Moravian- Campanian e Low diversity and abundance Not speci ed. Knobloch and Mai, 1991 Cretaceous Silesian Bezkydy Mts. Maastrichtian of angiosperm fruits & seeds. New Zealand South of Dunedin, Taratu Fm latest Maastrichtian Fluvial plane Angiosperm fruits, seeds Lignitised angiospe rms, Cantrill et al., 2011 Kaitangata Coalfield. mudstones. & flowers (Aff. Lauraceae). conifer leaves, shoots,

scales & seeds. Resear Germany Walbeck Maastrichtian High abundance of angiosperm Not speciﬁed. Knobloch and Mai, 1991

fruits & seeds. Low diversity ch

of species including Theaeceae 36

& Cyrillaceae. (20 ﬁ

Germany Eisleben Maastrichtian High abundance of angiosperm Not speci ed. Knobloch and Mai, 1991 1 fruits & seeds. Low diversity 2) 1 of species including Theaeceae 62 e

& Cyrillaceae. 1 9

Austria Kossen Niedendorf Gosau Fm Maastrichtian High abundance of angiosperm Not specified. Knobloch and Mai, 1991 0 fruits & seeds including Cyrillaceae affinities. Nigeria Mamu Fm Maastrichtian Coal. Inertinite (mmf) Average of 4 samples 17.5%, 10e20% reported in Glasspool and Scott (2010) U.S.A. Colorado, Denver Laramie Fm Maastrichtian Coal. Inertinite (mmf) Data from Glasspool Coalfield 20.7%, >20% and Scott (2010) U.S.A. Alabama McNairy Fm Maastrichtian Coal. Inertinite (mmf) Data from Glasspool 24.8%, >20% and Scott (2010) Spain Pedraforca, Maastrichtian Coal. Inertinite (mmf) Data from Glasspool Saldes Basin 39.2%, >20% and Scott (2010) Mexico Fuentes - Rio Maastrichtian Coal. Inertinite (mmf) Data from Glasspool Escondido Basin 19.1%, 10e20% and Scott (2010) U.S.A. New Mexico Sugarite Coal KeT boundary Coal. Inertinite (mmf) Data from Glasspool 27.9%, >20% and Scott (2010) 1 7 5 176 S.A.E. Brown et al. / Cretaceous Research 36 (2012) 162e190

Fig. 4. Charred plants from the Wealden of Shepherd’s Chine, Isle of Wight, England. A, siltstone with charred ferns, mainly Weichselia pinnules; FMNH PP55329 (Field Museum of Natural History, Chicago). B, detail of A showing charred fern pinnules of Weichselia and Phlebopteris. C, detail of A showing charred Weichselia pinnules. D, thin section of siltstone runnel showing layers with charred fern pinnules; FMNH PP55330. E, thin section of siltstone with concentrated layers of charred fern pinnules; FMNH PP55331. F, coarse sandstone with conifer wood charcoal; FMNH PP55332. G, thin section of F showing conifer wood charcoal; FMNH PP55332.

(so long as the biome continues to exist)” (p.19). This new concept is (e.g., those in Fig. 3) should form the nucleus for beginning to define of interest in geological studies, although interpretations of fire size super fire regimes where they can be linked to climatic and biome and frequency may be difficult to quantify (see for example data. Hudspith et al., 2012). Through the Cretaceous the nature of the A fire system is a concept that embraces the range of fire regimes vegetation affected by fire changed (section 3 above). These data at a given point in geological time. Fire systems are constrained to S.A.E. Brown et al. / Cretaceous Research 36 (2012) 162e190 177

Fig. 5. Fern-dominated charcoal from the Wealden of Beare Green, Surrey, UK (details in Harris, 1981). Scales in mm. A, block of siltstone with abundant charred fern pinnules; Natural History Museum, London; NHMUK V.60444. B, detail of another block showing charred pinnules of Weichselia and Phlebopteris; NHMUK V.60435. a particular time interval as they encapsulate not only temporally 4.2. Geographic distribution of fires defined vegetation types but also a specific range of atmospheric and climatic conditions. Throughout the Phanerozoic atmospheric The charcoal data reviewed in section 3 (Tables 1e3) and plotted oxygen concentrations have fluctuated dramatically (Bergman in Fig. 3 demonstrate widespread occurrences of fire across many et al., 2004; Berner, 2009; Glasspool and Scott, 2010) (Fig. 1). regions in the Cretaceous. A paucity of records in the Southern These perturbations will have affected fire regimes: during periods Hemisphere could reflect a lack of charcoal research and recogni- when atmospheric oxygen was elevated, climate will have exerted tion or the predominant occurrence of wildfires in the Northern a less significant control upon the distribution of fire than during Hemisphere (Bond and Scott, 2010). Clearly, future research at periods when it was lower. During the Cretaceous each of the Southern Hemisphere sites would benefit our understanding of any variables, vegetation, climate and atmospheric oxygen concentra- link between fire occurrence, Cretaceous palaeogeography and tion changed greatly, making an analysis of fire regimes and fire climates. Most studies on Cretaceous vegetation and climate (e.g., systems a significant challenge. Horrell, 1991; Spicer and Corfield, 1992; Coiffard et al., 2006, 2007;

Fig. 6. Fern-dominated charcoal from the Wealden of Hautrage, Belgium, (see Gomez et al., 2008, in press, for more information). A, reworked litter layer block (rip-up clast) from sandstone channel composed of two fern-dominated charcoal layers with clay-rich layer between; FMNH PP55333. B, fern rachis TS from polished block of specimen shown in A. C, macro- and meso-charcoal fragments macerated from the charcoal layers illustrated in A. This assemblage comprises predominantly ferns illustrated in Fig. 7. 178 S.A.E. Brown et al. / Cretaceous Research 36 (2012) 162e190 S.A.E. Brown et al. / Cretaceous Research 36 (2012) 162e190 179

Sellwood and Valdes, 2006; Sewall et al., 2007; Hay, 2008; of traits that allow them to cope with fire and even in some cases Donnadieu et al., 2009; Herman and Spicer, 2010) have not yet require fire (Pausas and Keeley, 2009; Bond and Scott, 2010; Keeley incorporated fire as a consideration. This is clearly a relevant factor et al., 2011; Crisp et al., 2011). The ability of a plant to survive fire across many areas (Fig. 3). obviously conveys a competitive advantage to a plant in a high-fire Though collection bias must be considered, a pattern emerges world. We note that the phenomenon of re-sprouting has been when charcoal mesofossil data are plotted on time-sliced palae- considered as an important aspect of fire survival and it is found in ogeographic maps (Fig. 3). A large number of charred angiosperm many plant groups that have lineages back to the Cretaceous reproductive organs occur from the Early and early Late Cretaceous (Pausas and Keeley, 2009; Crisp et al., 2011). Tree bark has also been of Western Europe and eastern North America (Tables 2, 3, Fig. 3). cited as being linked to surviving fire, including that of Pinus (see However, there are as yet no records from western North America, He et al., 2012; Belcher et al., in press for discussion), playing an despite extensive evidence of fire as indicated by inertinite in coal important role in many fire-prone ecosystems today and whose (Fig. 3). This suggests a search for mesofossil assemblages might be radiation may be rooted in the Cretaceous (Miller, 1999; He et al., productive. Equally the occurrence of charcoal in some African 2012). Serotony is another trait widely linked to fire and in the coals (Wuyep and Obaje, 2010) also indicates that the absence of Cretaceous charred serotinous cones occur in the earliest Creta- mesofossil records is not related to an absence of fire in the system. ceous (Pausas and Keeley, 2009) as do sclerotic closed fern indusia that may have been dropped or opened following fire (Watson and 4.3. Atmospheric oxygen content and a Cretaceous “high-fire” Alvin, 1996; Allen, 1998; Pausas and Keeley, 2009) (Fig. 7G). world Brentnall et al. (2005) compared data from the Cretaceous of Svalbard (Arctic) and Alexander Island in the Antarctic and showed There has been much debate over the atmospheric O2 levels that the conifers exhibited differences in their leaf lifespan. In required to allow combustion. This expansive debate is summarised combination with climate models these data were used to suggest elsewhere (e.g., Wildman et al., 2004; Belcher and McElwain, 2008; that fire was a critical factor in promoting the growth of trees with Belcher et al., 2010b). However, from the most recent experimen- short leaf lifespans, such as those found in the Arctic regions tation (Belcher et al., 2010b) it appears that for large wildfires to (Brentnall et al., 2005). occur atmospheric oxygen must exceed 17%. Fire frequency may also control the nature of the vegetation, Models have varied in their predictions of Cretaceous atmo- preventing the normal climax vegetation from developing (Bond spheric oxygen concentration (Berner et al., 2003; Bergman et al., et al., 2005). Herbaceous vegetation may allow rapid cool surface 2004; Berner, 2006, 2009; Glasspool and Scott, 2010) (Fig. 1). fires to burn. In such cases the fire effect on the soil may be minimal Berner (2009) predicted levels below present until the Albian, and the roots of the plant may not be killed, allowing for rapid re- rising just above 21% thereafter. Bergman et al. (2004) predicted growth following rain. If fires are frequent then the fire may kill any levels significantly above present throughout the Cretaceous but sapling of a larger tree, so that the herbaceous vegetation is reaching their highest during the Cenomanian. Similarly, Glasspool maintained (Bond and Keeley, 2005). The development of a forest and Scott (2010) predicted levels above present throughout the may change the local climate and reduce fire frequency. The Cretaceous but more moderately so and peaking at about the herbaceous plantefire cycle is one likely to be favoured during AlbianeCenomanian transition. Despite differences of degree, all times of warmth and high CO2 and high O2 (see Bond and Scott, three models predict a roughly similar pattern for Cretaceous 2010 for a review of this topic). atmospheric oxygen levels: falling initially from the Jurassic The large scale burning of woody vegetation will also have through the Berriasian then rising to a peak above present day a major impact on the carbon cycle (Fig. 12; Berner et al., 2003). levels somewhere near the Cenomanian before gradually declining Charcoal is a major residue of wildfire that because of its inert (Fig. 1). nature and post-fire erosion may be preferentially buried. This Under high oxygen conditions precipitation will be less of creates a positive feedback that produces further oxygen and a control on fire occurrence and fire will be less of a marker of increases fire activity (see Berner et al., 2003 for detailed discussion aridity (as indicated by Finklestein et al., 2005). Higher oxygen of this feedback; Fig. 12). would allow much wetter vegetation to burn than at the present The widespread record of fire as indicated here (Fig. 3) shows day. The review in section 3 (Tables 1e3) documents a large that the Cretaceous is, therefore, an important period of geological number of charcoalified plant mesofossil assemblages from the time with regard to a range of feedback mechanisms that may have Valanginian to the Campanian when oxygen levels are considered an impact on the Earth system. It may be difficult to jump from to be at their height (Fig. 1). a local scale of fire occurrence to a global scale of fire and its In combination, the records of charcoal in coal (Glasspool and significance, yet the widespread occurrence and record of fire in the Scott, 2010), the large numbers of charcoalified plant mesofossil Cretaceous (section 3; Fig. 3) and the inferred high atmospheric assemblages (section 3) and the inferred high atmospheric oxygen oxygen levels that would promote fire, lead to the conclusion that levels suggest a high-fire world during the Cretaceous and espe- such a jump in scale is important. These feedbacks indicate wide- cially in the late Early and early to middle Late Cretaceous. spread linkages such as charcoal production, erosion, deposition, elemental transport and carbon burial, all of which have impacts far 4.4. Vegetation and fire: forcings and feedbacks beyond the local area of a single fire. Amongst various models of Cretaceous climate and vegetation Fire may be influenced by and influence the nature of the (e.g., Wolfe and Upchurch, 1987; Otto-Bliesner and Upchurch, 1997; vegetation itself (Bond et al., 2005). Plants have evolved a number Haywood et al., 2004; Sellwood and Valdes, 2006), only a few have

Fig. 7. Scanning electron micrographs (SEMs) of charred conifers and ferns from the Wealden of Hautrage, Belgium (sample illustrated in Fig. 6A) (see Gomez et al., 2008, in press for more information). A, pinnule, probably of the fern Gleichenites; FMNH PP55334. B, detail of A showing stomata. C, scale, probably from the fern Gleichenites; FMNH PP55335. D, pinnule of the fern Phlebopteris showing the position of sori; FMNH PP55336. E, detail of D showing position of sorus and surface structure; F, pinnule of the fern Weichselia; FMNH PP55337. G, fern indusium, probably Weichselia; FMNH PP55338. H, fern rachis with pinnule attachment; FMNH PP55339. I, detail of H showing probable location of sorus. J, probable fern frond fragment; FMNH PP55340. K, detail of J showing three-dimensional anatomical preservation. L, conifer leafy shoot; FMNH PP55341. M, detail of L showing stomata. N, arthropod coprolite; FMNH PP5534; O, fragment of arthropod, probably also charred; FMNH PP55343. 180 S.A.E. Brown et al. / Cretaceous Research 36 (2012) 162e190

Fig. 8. SEMs of wood from the Coniacian Kamikituba assemblage, Japan (see Takahashi et al., 1999a, b; 2008 for more information). A, B, charred conifer wood charcoal. A, transverse section; B, detail of tracheids in A showing homogenized cell walls; FMNH PP55344. C, D, charred angiosperm wood. C, transverse section; D, tangential longitudinal section showing vessel end plate adjacent to ray at left; FMNH PP55345.

included fire feedbacks, e.g., Brentnall et al. (2005) for climate and to a reduction in evapo-transpiration, interception and surface Bergman et al. (2004) for atmospheres. Given the importance of storage for rainfall, resulting in greater levels of surface runoff and fires in the Cretaceous, demonstrated by the review in section 3, erosion (Shakesby and Doerr, 2006). Cannon et al. (2010, p.128) and the inference of a Cretaceous ‘high-fire’ world (section 4.3) it is indicated that continuous runoff can lead to “rapid and pervasive hoped that modellers will be convinced of the importance of fire overland flow”. feedbacks and incorporate them in future models. Shakesby and Doerr (2006) have calculated that the amount of ground covered by vegetation and litter is strongly linked to over- 4.5. Post-fire erosion land flow amounts after a rainfall event, reporting that for ground coverage of 37% only “14% of rainfall contributes to overland 4.5.1. Causes and effects of post-fire erosion flow”(p.275). However if only 10% ground coverage is present Wildfires and associated post-fire erosion can have a consider- a much higher percentage of rainfall contributes to overland flow, able effect on the hydrological cycle of a watershed (Moody and approximately “73% of rainfall” (Shakesby and Doerr, 2006). These Martin, 2009; Cannon et al., 2010 ) and can dramatically alter the figures clearly show how overland flow could be dramatically geomorphology of landscapes (Shakesby and Doerr, 2006). This increased by a wildfire, particularly a surface fire with a high erosional process includes the detachment, transport and deposi- severity. tion of sediment particles due to the influencing factors of both Intense heating of the soil reduces the stability of soil structure water and gravitational forces (Moody et al., 2008, Fig. 13). The and enhances the chances of erosional processes affecting the soil effect of fire upon soil profiles, vegetation coverage and bedrock all profile. Ash and charcoal produced by fire can seal soil macropores contribute to increasing susceptibility to enhanced erosion. This in (Nyman et al., 2010) reducing infiltration levels further. Soil turn can lead to extensive sediment transport and deposition hydrophobicity is acknowledged as one of the dominant factors (Moody and Martin, 2009, Fig. 13). influencing accelerated slope runoff and erosional levels after a fire Wildfires can result in the burning and destruction of both event (Shakesby and Doerr, 2006; Doerr et al., 2009; Beatty and vegetation and litter layers, which can have a dramatic effect on the Smith, 2010; Finley and Glenn, 2010). In addition to water repel- hydrological cycle, altering many components (Shakesby and Doerr, lent compounds coating soil particles during a wildfire, organic 2006). This partial or full removal of vegetation and litter can lead substances, which are hydrophobic in nature, already located S.A.E. Brown et al. / Cretaceous Research 36 (2012) 162e190 181

Fig. 9. SEMs of charred plant mesofossils from the Late Cretaceous (Santonian) Gaillard Formation of Allon, Georgia, USA (see Herendeen et al., 1999 for descriptions of taxa). A, Antiquacupula sulcata Sims, Herendeen and Crane, staminate ﬂower originally with six tepals arranged in two cycles; FMNH PP55041. B, Antiquacupula sulcata, staminate ﬂower originally with six tepals arranged in two cycles; FMNH PP55042. C, adaxial surface of fern pinnule; FMNH PP55043. D, adaxial surface of fern with alternate pinnules; FMNH PP45162. E, moss, gametophytic stage; FMNH PP55044. F, Eopolytrichum antiquum Konopka, Herendeen, Smith, Merrill and Crane, sporophyte capsule; FMNH PP44717.

within both the litter layer and upper soil profile are volatilised 2010) of water repellent soil properties, which makes the erosion (Shakesby and Doerr, 2006). This volatilisation creates a “pressure levels variable in a burned area. gradient” within the area heated, driving some compounds into the Both reduced soil stability and increased soil hydrophobicity atmosphere and others into lower soil layers where they condense result in reduced infiltration levels through the soil, and thus (Shakesby and Doerr, 2006). The heat derived from a wildfire will enhance overland flow/runoff levels leading to erosion. Soil both create and redistribute already present hydrophobic material. degradation, involving the creation of rills, can be enhanced via This redistribution results in “spatial variability” (Nyman et al., increased overland flow due to limited infiltration. 182 S.A.E. Brown et al. / Cretaceous Research 36 (2012) 162e190

Fig. 10. Charcoal from the Upper Cretaceous of Scania, Sweden. A, thin section of sediment showing laminations and ﬁne charcoal layers. B, SEM of charred Scandianthus ﬂower (see Friis and Skarby, 1981) (illustration after Scott, 2000 from specimen supplied by E.M. Friis).

The combination of rock weathering, vegetation and litter a post-fire erosion deposit. Other probable post-fire erosion- removal and the creation of a hydrophobic soil layer all contribute deposition sediments from the Isle of Wight include sheet-flood to increased surface runoff and overland flow. It is this surface plant-debris beds containing charcoal and vertebrates movement of water that results in transportation and deposition of (Sweetman and Insole, 2010). In the Wealden of Hautrage in large sediment loads that may occur within, and down-channel Belgium alternating layers of fern charcoal and clay may repre- from, the burned area (Cannon et al., 2010). The degree to which sent post-fire runoff into quiescent waters (Fig. 6A). These a watershed is affected by post-fire erosion is controlled by many sediments are also associated with fluvial sands containing factors, as outlined by Moody and Martin (2001, 2009), including abundant charcoal (Gomez et al., 2008) that are themselves fire severity, prior sensitivity of the affected area to erosion, probably fire related. Scott and Stea (2002) noted quartz grains precipitation and the frequency of wildfires. One of the most amongst lignites with charcoal and these also may reflect important factors is the amount of precipitation, as without suffi- evidence of post-fire erosion. cient rainfall after the wildfire event post-fire erosion will not New data on Campanian sequences in Dinosaur Provincial occur. Park, Alberta, indicate that significant post-fire erosion occurred within that system. The area was fluvially active during the 4.5.2. The potential role of post-fire erosion in the Cretaceous Campanian (Currie and Koppelhus, 2005). In the single strati- As outlined above, post-fire erosion frequently follows fire. graphic profile analysed to date (by SAEB), dinosaur bone beds Although widely recognised following modern fires, its occur- containing disarticulated bones and some articulated bones, have rence may be overlooked in the geological record and there are been found to co-occur with charcoal deposits (Figs. 14 and 15). few reported examples of this phenomenon in the Cretaceous. The presence of these together on the same stratigraphic horizon The coarse sands from the Wealden of the Isle of Wight is hypothesised to represent post-fire erosion deposition whereby (Collinson et al., 2000; Scott, 2000) are associated with abundant flooding, as a result of rainfall after fire events, has washed both charcoal and this chaotic mix (Fig. 4F) probably represents dinosaurs and charcoal into one deposit. This hypothesis is being

Fire Severity Fire Ecosystem or Intensity response burn severity

Energy released Organic matter loss Erosion/ Vegetational recovery

Fig. 11. The relationship of ﬁre intensity, severity and ecosystem response (after Keeley, 2009). S.A.E. Brown et al. / Cretaceous Research 36 (2012) 162e190 183

FIRE

Erosion

Charcoal O2 Land Plants

Organic C sed. Burial

Fig. 12. Systems diagram of fire and oxygen feedbacks (after Berner et al., 2003). Systems analysis showing the feedbacks between fire and atmospheric oxygen: arrows originate with causes and end at effects. Straight arrows lead to a positive response (e.g., oxygen increases, fires increase) and arrows with bull’s-eyes are negative responses (e.g., fires increase, vegetation decreases). Closed loops with an even number of bull’s-eyed arrows or solely plain arrows are positive feedbacks, and those with an odd number of arrows with bull’s eyes are negative feedbacks (see Berner et al., 2003 for further discussion).

tested by study of additional sites and sequences (SAEB in prog- Dinosaur bone beds (containing Albertosaurus) in the Maas- ress). While we do not know of any other Cretaceous dinosaur trichtian part of the sequence in Dry Island National Park around accumulations that have been directly linked to fire events, such Drumheller (Alberta) also occur associated with rapidly deposited vertebrate accumulations with charcoal have been reported in the fluvial deposits (Eberth and Currie, 2010). We suggest that such Cretaceous (Tables 1e3) and elsewhere in the fossil record (e.g., deposits be searched for charcoal sothat their possible origin as a post- Sander, 1987). fire erosion deposit can be evaluated. The full effects of wildfire on

Fig. 13. A, B, charcoal-rich flows after a forest fire by overland flow following a rainstorm: the Rodeo-Chediski Fire, Apache-Sitgreaves National Forest, Arizona, USA, 2002 (Photographs courtesy of D. Neary, US Geological Survey). 184 S.A.E. Brown et al. / Cretaceous Research 36 (2012) 162e190

Fig. 14. Charcoal from the Campanian sediments of Dinosaur Provincial Park, Alberta, Canada. Specimens to be deposited in the Royal Tyrrell Museum, Drumheller, Canada. A, panoramic view of sediments in Dinosaur Provincial Park. B, C, fluvial sandstone with charcoal (arrows). D, large fragments of gymnosperm wood charcoal extracted from sandstone unit. E, small charcoal fragments extracted from siltstone unit. F, SEM of transverse section of charred gymnosperm wood showing distinctive growth rings. G, SEM of tangential longitudinal section of charred gymnosperm wood showing uniseriate rays and a small number of bordered pits. H, SEM of radial longitudinal section of charred gymnosperm wood showing simple oblique cross field pitting in the rays. vegetation, litter, soil horizons and lithologies need to be considered Cretaceous (Niklas et al., 1985; Crane and Lidgard, 1989; Lidgard when analysing ancient fires, along with the role of post-fire erosion. and Crane, 1990; Lupia et al., 1999; Friis et al., 2010) and by the Late Cretaceous had risen to dominance (Lidgard and Crane, 1990; 4.6. Fire and the Cretaceous spread of angiosperms Wing and Boucher, 1998; Lupia et al., 1999; Friis et al., 2006) (Fig. 1). Clearly the nature of world vegetation and biome composition The fossil record demonstrates that angiosperms became an changes rapidly through the period (Saward, 1992; Wing and increasingly significant component of the flora through the Boucher, 1998; Spicer, 2003). While angiosperms may have had S.A.E. Brown et al. / Cretaceous Research 36 (2012) 162e190 185

Fig. 15. Example lithological log of a small sedimentary unit in Dinosaur provincial Park, Alberta, Canada showing numerous charcoal-bearing horizons and the position of dinosaur remains. The charcoal shown on this sedimentary log was visible in the field with the naked eye. The large amount of charred plant material highlights the importance of wildfire as an ecosystem process during the Late Campanian in Dinosaur Provincial Park, likely to have had environmental impact on both the local vegetation and the dinosaurs inhabiting the landscape at this time. The multiple charcoal horizons indicate that wildfire was a typical environmental factor during the 1.7 myr represented by deposits exposed within the park. It can be hypothesised that both the fire itself and associated post-fire erosion would have impacted on the life of the dinosaurs through vegetation loss and potential flash flood.

a competitive advantage over gymnosperms, Bond and Scott (2010) Little angiosperm wood has yet been identified at a time when suggested that a high oxygen world during the Cretaceous may the earliest flowering plants first appear in the fossil record, despite have aided the diversification and spread of these plants through the occurrence of pollen, leaves and flowers (Friis et al., 2006). It has the promotion of wildfire activity. The fuller literature compilation been proposed that early flowering plants were essentially weedy presented here underlines the widespread association of fire and plants, generally small and herbaceous or, at most, shrubby with early angiosperms. little wood (Taylor and Hickey, 1996; Royer et al., 2010). The more 186 S.A.E. Brown et al. / Cretaceous Research 36 (2012) 162e190

impact of extensive fire is rarely considered. Fire in the short term may restrict food availability and promote animal movement. Those herbivores that favour lush growth of herbaceous plants would be aided by the development of an angiospermefire cycle (section 4.6). It has been speculated that, in the Carboniferous and Permian (Sander, 1987; Scott and Jones, 1994), fire may have driven animals to their death. The same is possible in the Cretaceous. Furthermore, during post-fire erosion (section 4.5) flooding and sediment movement can be very rapid indeed. If animals were feeding on vegetation near a stream then even a storm following a fire many tens of kilometres away may have had a major impact, causing not only the death of dinosaurs but also their entombment and preservation. The abundance of charcoal associated with dinosaur deposits in the Campanian of Dinosaur Provincial Park (section 4.5.2) holds out this scenario, but our research is at too early a stage to make a definite link.

4.8. An impact on the marine realm? Marine anoxia

Kump (1988) has hypothesised a relationship between fire, oxygen and phosphorous (Fig. 16). Post-fire erosion, especially fi Fig. 16. Systems diagram of re-phosphorous feedback cycle (after Kump, 1988). during periods of particularly high fire activity, would have flushed Positive feedbacks are shown by arrows and negative feedbacks are shown by bull’s- eyes. In this case increasing fire leads to increase in phosphorous to the ocean allowing phosphorous into the oceans. Whether this would have resulted in the rapid growth of plankton that in turn promotes widespread anoxia and marine elevated algal productivity and subsequent marine anoxia is more carbon burial. speculative. However, it may be a significant short-term feedback mechanism, though Lenton and Watson (2000) suggested that in the long term increases in oxygen are counteracted by reductions in phosphorous weathering flux and organic carbon burial and via comprehensive literature compilation presented here reinforces versa. fi the view that the earliest angiosperms subjected to re had little The relationship between nutrient cycling, phosphorous and wood. Studies on the ecology of these early angiosperms suggest the widespread anoxia leading to the deposition of organic-rich they were well adapted to disturbed environments, early forms shales (Oceanic Anoxic Event: OAE) has been discussed by living near active stream systems or wet to fully aquatic environ- Berrocoso et al. (2010) and Kraal et al. (2010). In particular, the ments (Doyle and Hickey, 1976; Hickey and Doyle, 1977; Friis et al., CenomanianeSantonian OAE coincides with an inferred Creta- fi 2010). Sapindopsis may represent an example of a post- re recovery ceous oxygen high (Glasspool and Scott, 2010) (Fig. 1) while species from this time interval (Taylor and Hickey, 1996), its leaves contemporaneous coals in West African coals (Wuyep et al., 2010) frequently being found above horizons full of charcoal (Hickey and have high charcoal contents that indicate a period of increased fire Doyle, 1977). activity (Fig. 3). While it is clear that not all OAEs are necessarily Angiosperms possessed innovations enabling rapid growth linked to fire it may have been a contributing factor at least in fi (Brodribb and Field, 2010). For example, ef cient hydraulic systems some cases. A search for fire-derived components in such black associated with high leaf venation densities (Boyce et al., 2009) and shales would be worthwhile to establish the importance of wild- large leaf areas would have allowed increased uptake of CO2 fires in such events. (Brodribb and Field, 2010), especially during a time interval (Late Cretaceous) when CO2 was dropping (Fletcher et al., 2008; Quan et al., 2009). This efficiency would have allowed a doubling of 5. Conclusions a maximum photosynthetic rate for Late Cretaceous angiosperms fi and so led to accelerated fuel accumulation. This build-up of fuel This literature compilation of Cretaceous charcoali ed plant would have provided a feedback response promoting burning and mesofossils shows that charcoal is widely distributed throughout fi the development of a weedy angiospermefire cycle (Bond and the Cretaceous, indicating the extensive occurrence of wild re. Scott, 2010). The inferences (section 4.3) that higher Cretaceous Records are concentrated in the Northern Hemisphere but this oxygen levels would have promoted fire within wetter vegetation is most likely a result of either a lack of appropriate sedimentary than would burn in the modern world and that rapid fire return settings or limited research focused on charcoals. Results intervals would prevent the growth of competing trees are also from ongoing work (SAEB) on charcoals from the Campanian of important. Therefore, the establishment of an angiospermefire Dinosaur Provincial Park in Canada show just how likely it is cycle may have been responsible, in the mid- to Late Cretaceous, that numerous charcoal-rich deposits have been previously for promoting the diversification and spread of angiosperms, as overlooked. fi suggested by Bond and Scott (2010). Charcoali ed mesofossils provide a wealth of plant organs and tissues with 3-D anatomical data that have contributed to our understanding of not only the evolution of Cretaceous flowering 4.7. Scenarios of impacts on herbivores plants but also the development of Cretaceous vegetation from a world without flowering plants to one in which they dominate. The Cretaceous was a period with large herbivores. The rela- Wildfire may have played a significant role in promoting the spread tionship between long-necked and short-necked herbivores and of weedy angiosperms that were well adapted to living in disturbed vegetation has long been discussed (Bakker, 1978). However, the environments. S.A.E. Brown et al. / Cretaceous Research 36 (2012) 162e190 187

In the modern Earth system wildfire is controlled in part by fuel Belcher, C.M., Collinson, M.E., Scott, A.C., 2005. Constraints on the thermal energy moisture, but this may be partly overridden in the Cretaceous released from the Chicxulub impactor: new evidence from multi-method charcoal analysis. Journal of the Geological Society, London 162, 591e602. where it is suggested that the atmospheric oxygen concentration Belcher, C.M., Finch, P., Collinson, M.E., Scott, A.C., Grassineau, N., 2009. Geochem- was greater, perhaps over 25%. This would enable wetter vegetation ical evidence for combustion of hydrocarbons during the K-T impact event. to burn and allow more frequent and more widespread fires. Fires Proceedings of the National Academy of Sciences, USA 106, 4112e4117. fi Belcher, C.M., Mander, L., Rein, G., Jervis, F.X., Haworth, M., Hesselbo, S.P., may have played a particularly signi cant role in the mid- Glasspool, I.J., McElwain, J.C., 2010a. Increased fire activity at the Triassic/ Cretaceous as angiosperms began to diversify and oxygen levels Jurassic boundary in Greenland due to climate-driven floral change. Nature were at their highest. Geoscience 3, 426e429. fi Belcher, C.M., Yearsley, J.M., Hadden, R.M., McElwain, J.C., Rein, G., 2010b. Baseline Post- re erosion is likely to have played a major role in Creta- intrinsic flammability of Earths’ ecosystems estimated from paleoatmospheric ceous terrestrial environments, may have been responsible for at oxygen over the past 350 million years. Proceedings of the National Academy of least some dinosaur bone bed deposits and may have added to Sciences, USA 107, 22448e22453. Belcher, C.M., Collinson, M.E., Scott, A.C. A 400 million year record of fire. In: phosphorous runoff into oceanic settings contributing to anoxia. Belcher, C.M., Rein, G. (Eds), Fire Phenomena in the Earth System e An Inter- Abundant wildfire events, producing abundant charcoal, will have disciplinary Approach to Fire Science. J. Wiley and Sons, Chichester, in press. had a significant effect upon the sedimentary carbon cycle. Bergman, N.M., Lenton, T.M., Watson, A.J., 2004. COPSE: a new model of biogeochemical e Although recent evidence suggests that extensive wildfires were cycling over Phanerozoic time. American Journal of Science 304, 397 437. Berner, R.A., 2006. GEOCARBSULF: a combined model for Phanerozoic atmospheric fi not a factor in CretaceousePaleogene boundary events, wild re is O2 and CO2. Geochimica et Cosmochimica Acta 70, 5653e5664. shown here to be an important part of the Cretaceous Earth system Berner, R.A., 2009. Phanerozoic atmospheric oxygen: new results using the GEO- and should be included in coupled climate-vegetational models. CARBSULF model. American Journal of Science 309, 603e606. fi Berner, R.A., Beerling, D.J., Dudley, R., Robinson, J.M., Wildman, R.A. Jr., 2003. The Cretaceous is here judged to have been a “high- re” world and Phanerozoic atmospheric oxygen. Annual Review of Earth and Planetary further studies on Cretaceous charcoal assemblages are needed to Sciences 31, 105e134. unlock the palaeoenvironmental and palaeoecological information Berrocoso, A.J., MacLeod, K.G., Martin, E.E., Bourbon, E., Londoño, C.I., 2010. Nutrient trap for Late Cretaceous organic-rich shales in the tropical North Atlantic. in these important deposits. Geology 38, 1111e1114. Bond, W.J., Keeley, J.E., 2005. Fire as global ‘herbivore’: the ecology and evolution of flammable ecosystems. Trends in Ecology and Evolution 20, 387e394. Acknowledgements Bond, W.J., Scott, A.C., 2010. Fire and the spread of flowering plants in the Creta- ceous. New Phytologist 188, 1137e1150. Bond, W.J., Woodward, F.I., Midgley, G.F., 2005. The global distribution of ecosys- We thank Chris Scotese for providing the palaeogeographic tems in a world without fire. New Phytologist 165, 525e538. maps. We thank Dennis Braman for aiding ACS and SAEB in their Bowman, D.M.J.S., Balch, J.K., Artaxo, P., Bond, W.J., Carlson, J.M., Cochrane, M.A., fieldwork in Dinosaur Provincial Park. ACS acknowledges the help D’Antonio, C.M., DeFries, R.S., Doyle, J.C., Harrison, S.P., Johnston, F.H., of Leo Hickey, Scott Wing, John Calder, Ralph Stea, William Bond, Keeley, J.E., Krawchuk, M.A., Kull, C.A., Marston, J.B., Moritz, M.A., Prentice, I.C., Roos, C.I., Scott, A.C., Swetnam, T.W., van der Werf, G.R., Pyne, S.J., 2009. Fire in Johan Yans, Phillipe Gerienne, Gary Nichols and Masamichi Taka- the Earth System. Science 324, 481e484. hashi for help in the field and loan of specimens. ACS thanks Yale Boyce, C.K., Brodribb, T.J., Field, T.S., Zwieniecki, M.A., 2009. Angiosperm leaf vein University, Royal Holloway Research Strategy Fund and a personal evolution was physiologically and environmentally transformative. Proceedings e fi of the Royal Society, B 276, 1771 1776. charitable donation for supporting his research on Cretaceous res. Brasier, M., Cotton, L., Yenney, 2009. First report of amber with spider webs and We thank Sharon Gibbons, Neil Holloway and Kevin D’Souza for microbial inclusions from the earliest Cretaceous (c. 140Ma) of Hastings, Sussex. technical support. We thank Tim Ewin for arrangements and the Journal of the Geological Society, London 166, 989e997. Brentnall, S.J., Beerling, D.J., Osbourne, C.P., Harland, M., Francis, J.E., Valdes, P.J., Natural History Museum, London, photographic unit for photog- Wittig, V.E., 2005. Climatic and ecological determinants of leaf lifespan in polar raphy enabling us to include the images in Fig. 5. We thank David forests of the high CO2 Cretaceous ‘greenhouse’ world. Global Change Biology Batten and two anonymous referees for their constructive criticism, 11, 2177e2195. Brodribb, T.J., Field, T.S., 2010. Leaf hydraulic evolution led a surge in leaf photo- which much improved the paper. synthetic capacity during early angiosperm diversification. Ecology Letters 13, 175e183. Cannon, S.H., Gartner, J.E., Rupert, M.G., et al., 2010. Predicting the probability and References volume of postwildfire debris flows in the intermountain western United States. Geological Society of America. Bulletin 122, 127e144. Allen, P., 1998. PurbeckeWealden (Early Cretaceous) climates. Proceedings of the Cantrill, D.J., Wanntorp, L., Drinnan, A.N., 2011. Mesofossil flora from the Late Geologists’ Association 109, 197e236. Cretaceous of New Zealand. Cretaceous Research 32, 164e173. Alvin, K.L., 1974. Leaf anatomy of Weichselia based on fusainized material. Palae- Coiffard, C., Gomez, B., Kvacek, J., Thevenard, F., 2006. Early angiosperm ecology, ontology 17, 587e598. evidence from the AlbianeCenomanian of Europe. Annals of Botany 98, Appert, O., 2010. Die fossile Makroflora der Unterkreide (Hauterivian) von Manja in 495e502. Südwest-Madagaskar: Farne und Schachtelhalme (Filicatae und Equisetatae). Coiffard, C., Gomez, B., Thevenard, F., 2007. Early Cretaceous angiosperm invasion of Schweizerische Paläontologische Abhandlungen 129, 81 pp. western Europe and major environmental changes. Annals of Botany 100, Axsmith, B.J., 2006. The vegetative structure of a Lower Cretaceous conifer from 545e553. Arkansas: further implications for morphospecies concepts in the Cheir- Collinson, M.E., Featherstone, C., Cripps, J.A., Nichols, G.J., Scott, A.C., 2000. Charcoal- olepidiaceae. Cretaceous Research 27, 309e317. rich plant debris accumulations in the Lower Cretaceous of the Isle of Wight, Axsmith, B.J., Jacobs, B.F., 2005. The conifer Frenelopsis ramosissma (Cheirolepidaceae) in England. Acta Palaeobotanica Supplement 2, 93e105. the Lower Cretaceous of Texas: systematic, biogeographical and paleoecological Crane, P.R., Herendeen, P.S., 1996. Cretaceous floras containing angiosperm flowers implications. International Journal of Plant Sciences 166, 327e337. and fruits from eastern North America. Review of Palaeobotany and Palynology Bakker, R.T., 1978. Dinosaur feeding behaviour and the origin of flowering plants. 90, 319e337. Nature 274, 661e663. Crane, P.R., Lidgard, S., 1989. Angiosperm diversification and paleolatitudinal Batten, D.J., 1974. Wealden palaeoecology from the distribution of plant fossils. gradients in Cretaceous floristic diversity. Science 246, 675e678. Proceedings of the Geologists’ Association 85, 443e458. Crane, P.R., Upchurch, G.R., 1987. Drewria potomacensis gen. et sp. nov. American Batten, D.J., 1998. Palaeoenvironmental implications of plant, insect and other Journal of Botany 74, 1722e1736. organic-walled microfossils in the Weald Clay Formation (Lower Cretaceous) of Crane, P.R., Friis, E.M., Pedersen, K.R., 1989. Reproductive structure and function in southeast England. Cretaceous Research 19, 279e315. Cretaceous Chloranthaceae. Plant Systematics and Evolution 165, 211e226. Beatty, S.M., Smith, J.E., 2010. Fractional wettability and contact angle dynamics in Crane, P.R., Pedersen, K.R., Friis, E.M., Drinnan, A.N., 1993. Early Cretaceous (Early to burned water repellent soils. Journal of Hydrology 391, 97e108. Middle Albian) platanoid inflorescences associated with Sapindopsis leaves Belcher, C.M., 2009. Reigniting the Cretaceous-Palaeogene firestorm debate. from the Potomac Group of eastern North America. Systematic Botany 18, Geology 37, 1147e1148. 328e344. fl fi Belcher, C.M., McElwain, J.C., 2008. Limits for combustion in low O2 redefine Crepet, W.L., Nixon, K.C., 1992. Two new fossil owers of magnoliid af nity from the paleoatmospheric predictions for the Mesozoic. Science 321, 1197e1200. Late Cretaceous of New Jersey. American Journal of Botany 85, 1273e1288. Belcher, C.M., Collinson, M.E., Sweet, A.R., Hildebrand, A.R., Scott, A.C., 2003. Fireball Crisp, M.D., Burrows, G.E., Cook, L.G., Thornhill, A.H., Bowman, D.M.J.S., 2011. passes and nothing burns e the role of thermal radiation in the K/T event: Flammable biomes dominated by eucalypts originated at the Creta- evidence from the charcoal record of North America. Geology 31, 1061e1064. ceousePalaeogene boundary. Nature Communications 2, 1e8. 188 S.A.E. Brown et al. / Cretaceous Research 36 (2012) 162e190

Currie, P.J., Koppelhus, E.B., 2005. Dinosaur Provincial Park: a spectacular ancient Friis, E.M., Crane, P.R., Pedersen, K.R., 1997. Anacostia, a new basal angiosperm from ecosystem revealed. Indiana University Press, Bloomington, IN, 651 pp. the Early Cretaceous of North America and Portugal with trichotomocolpate/ Doerr, S.H., Woods, S.W., Martin, D.A., Casimiro, M., 2009. ‘Natural background’ soil monocolpate pollen. Grana 36, 225e244. water repellency in conifer forests of the north-western USA: its prediction and Friis, E.M., Pedersen, K.R., Crane, P.R., 1999. Early angiosperm diversification: the relationship to wildfire occurrence. Journal of Hydrology 371, 12e21. diversity of pollen associated with angiosperm reproductive structures in Early Donnadieu, Y., Goddéris, Y., Bouttes, N., 2009. Exploring the climatic impact of the Cretaceous floras from Portugal. Annals of the Missouri Botanical Garden 86, continental vegetation on the Mesozoic atmospheric CO2 and climate history. 259e296. Climate of the Past 5, 85e96. Friis, E.M., Pedersen, K.R., Crane, P.R., 2000. Fossil floral structures of a basal Doyle, J.A., Hickey, L.J., 1976. Pollen and leaves from the mid-Cretaceous Potomac angiosperm with monocolpate, reticulate-acolumellate pollen from the Early Group and their bearing on angiosperm evolution. In: Beck, C.B. (Ed.), Origin Cretaceous of Portugal. Grana 39, 226e239. and Early Evolution of Angiosperms. Columbia University Press, New York, NY, Friis, E.M., Pedersen, K.R., Schonenberger, J., 2003. Endressianthus, a new pp. 139e206. Normapolles-producing plant genus of fagalean affinity from the Late Creta- Drinnan, A.N., Crane, P.R., Friis, E.M., Pedersen, K.R., 1990. Lauraceous flowers from ceous of Portugal. International Journal of Plant Sciences 64 (5 Supplement), the Potomac Group (mid-Cretaceous) of eastern North America. Botanical 201e223. Gazette 151, 370e384. Friis, E.M., Pedersen, K.R., Crane, P.R., 2004. Araceae from the Early Cretaceous of Drinnan, A.N., Crane, P.R., Pedersen, K.R., Friis, E.M., 1991. Angiosperm flowers and Portugal: evidence on the emergence of monocotyledons. Proceedings of the tricolpate pollen of buxaceous affinity from the Potomac Group (mid-Creta- National Academy of Sciences, USA 101, 16565e16570. ceous) of eastern North America. American Journal of Botany 78, 153e176. Friis, E.M., Pedersen, K.R., Crane, P.R., 2006. Cretaceous angiosperm flowers: inno- Eberth, D.A., Currie, P.J., 2010. Stratigraphy, sedimentology, and taphonomy of the vation and evolution in plant reproduction. Palaeogeography, Palae- Albertosaurus bonebed (upper Horseshoe Canyon Formation; Maastrichtian), oclimatology, Palaeoecology 232, 251e293. southern Alberta, Canada. Canadian Journal of Earth Sciences 47, 1119e1143. Friis, E.M., Pedersen, K.R., Crane, P.R., 2010. Cretaceous diversification of angio- Eklund, H., 2003. First Cretaceous flowers from Antarctica. Review of Palaeobotany sperms in the western part of the Iberian Peninsula. Review of Palaeobotany and Palynology 127, 187e217. and Palynology 162, 341e361. Eklund, H., Kvacek, J., 1998. Lauraceous inflorescences and flowers from the Cen- Friis, E.M., Crane, P.R., Pedersen, K.R., 2011. Early flowers and angiosperm evolution. omanian of Bohemia (Czech Republic, central Europe). International Journal of Cambridge University Press, Cambridge, 596 pp. Plant Sciences 159, 668e686. Frumin, S.I., Friis, E.M., 1996. Liriodendroid seeds from the Late Cretaceous of Eklund, H., Cantrill, D.J., Francis, J.E., 2004. Late Cretaceous plant mesofossils from Kazakhstan and North Carolina, USA. Review of Palaeobotany and Palynology Table Nunatak, Antarctica. Cretaceous Research 25, 211e228. 94, 39e55. Endress, P.K., Friis, E.M., 1991. Archamamelis, hamamelidalean flowers from the Gandolfo, M.A., Nixon, K.C., Crepet, W.L., 1998. A new fossil flower from the Turo- Upper Cretaceous of Sweden. Plant Systematics and Evolution 175, 101e114. nian of New Jersey: Dressiantha bicarpellata gen. et sp. nov. (Cappareles). Eriksson, O., Friis, E.M., Pedersen, K.R., Crane, P.R., 2000. Seed size and dispersal American Journal of Botany 85, 964e974. systems of Early Cretaceous angiosperms from Famalicão, Portugal. Interna- Gandolfo, M.A., Nixon, K.C., Crepet, W.L., 2002. Triuridaceae fossil flowers from the tional Journal of Plant Sciences 161, 319e329. Upper Cretaceous of New Jersey. American Journal of Botany 89, 1940e1957. Falcon-Lang, H.J., Kvacek, J., Ulicny, D., 2001. Fire-prone plant communities and Gill, A.M., 1975. Fire and the Australian flora: a review. Australian Forestry 38, 4e25. palaeoclimate of a Late Cretaceous fluvial to estuarine environment, Pecínov Glasspool, I.J., Scott, A.C., 2010. Phanerozoic concentrations of atmospheric oxygen quarry, Czech Republic. Geological Magazine 138, 563e576. reconstructed from sedimentary charcoal. Nature Geoscience 3, 627e630. Falcon-Lang, H.J., MacRae, R.A., Csank, A.Z., 2004. Palaeoecology of Late Cretaceous Glasspool, I.J., Scott, A.C. Identifying past fire events. In: Belcher, C.M., Rein, G. (Eds), polar vegetation preserved in the Hansen Point Volcanics, NW Ellesmere Island, Fire Phenomena in the Earth System e An Interdisciplinary Approach to Fire Canada. Palaeogeography, Palaeoclimatology, Palaeoecology 212, 45e64. Science. J. Wiley and Sons, Chichester, in press. Falcon-Lang, H.J., Fensome, R.A., Gibling, M.R., Malcome, J., Flecher, K.R., Goldin, T.J., Melosh, H.J., 2009. Self-shielding of thermal radiation by Chicxulub Holleman, M., 2007. Karst-related outliers of the Cretaceous Chaswood impact ejecta: firestorm or fizzle? Geology 37, 1135e1138. Formation of Maritime Canada. Canadian Journal of Earth Sciences 44, Gomez, B., Martínez-Delclòs, X., Bamford, M., Philippe, M., 2002. Taphonomy and 619e642. palaeoecology of plant remains from the oldest African Early Cretaceous amber Finklestein, D.B., Pratt, L.M., Curtin, T.M., Brassell, S.C., 2005. Wildfires and seasonal locality. Lethaia 35, 300e308. aridity recorded in Late Cretaceous strata from south-eastern Arizona, USA. Gomez, B., Gillot, T., Daviero-Gomez, V., Spagna, P., Yans, J., 2008. Paleoflora from Sedimentology 52, 587e599. Wealden facies strata of Belgium: mega-and meso-fossils of Hautrage (Mons Finley, C.D., Glenn, N.F., 2010. Fire and vegetation type effects on soil hydrophobicity Basin). Memoirs of the Belgian Geological Survey 55, 53e60. and infiltration in the sagebrush-steppe: II. Hyperspectral analysis. Journal of Gomez, B., Gillot, T., Daviero-Gomez, V., Coiffard, C., Spagna, P., Yans, J. Chapter 10. Arid Environments 74, 660e666. Mesofossil plant remains from the BarremianeAptian of Hautrage (Mons Basin, Fletcher, B.J., Brentnall, S.J., Anderson, C.W., Berner, R.A., Beerling, D.J., 2008. Belgium) with taphonomy, palaeoecology and palaeoenvironment insights. In: Atmospheric carbon dioxide linked with Mesozoic and early Cenozoic climate Godefroit, P. (Ed.), Bernissart: Dinosaurs in Depth. Indiana University Press, change. Nature Geoscience 1, 43e48. Bloomington, IN, in press. Francis, J.E., 1983. The dominant conifer of the Jurassic Purbeck Fm, England. Grocke, D.R., Price, G.D., Robinson, S.A., Baraboshkin, E.Y., Mutterlose, J., Ruffell, A.H., Palaeontology 26, 277e294. 2005. The Upper Valanginian (Early Cretaceous) positive carbon-isotope event Francis, J.E., Coffin, M.F., 1992. Cretaceous fossil wood from the Raggan Basin, recorded in terrestrial plants. Earth and Planetary Science Letters 240, 495e509. southern Karguelen Plateau (site 750). Proceedings of the ODP Scientific Results Harris, T.M., 1981. Burnt ferns from the English Wealden. Proceedings of the 120, 273e280. Geologists’ Association 92, 47e58. Friis, E.M., 1983. Upper Cretaceous (Senonian) floral structures of juglandalean Harrison, S.P., Marlon, J., Bartlein, P.J., 2010. Fire in the Earth system. In: Dodson, J. affinity containing Normapolles pollen. Review of Palaeobotany and Palynology (Ed.), Changing Climates, Earth Systems and Society. Springer-Verlag, Berlin, 39, 161e188. pp. 21e48. Friis, E.M., 1984. Preliminary report of Upper Cetaceous angiosperm reproductive Harvey, M.C., Brassell, S.C., Belcher, C.M., Montanari, A., 2008. Combustion of fossil organs from Sweden and their level of organization. Annals of the Missouri organic matter at the K-P boundary. Geology 36, 355e358. Botanical Garden 71, 403e418. Haworth, M., Hesselbo, S.P., McElwain, J.C., Robinson, S.A., Brunt, J.W., 2005. Mid- Friis, E.M., 1985. Actinocalyx gen. nov., sympetalous angiosperm flowers from the Cretaceous pCO2 based on stomata of the extinct conifer Pseudofrenelopsis Upper Cretaceous of southern Sweden. Review of Palaeobotany and Palynology (Cheirolepidiaceae). Geology 33, 749e752. 45, 171e183. Hay, W.W., 2008. Evolving ideas about the Cretaceous climate and ocean circula- Friis, E.M., Pedersen, K.R., 1990. Structure of the Lower Cretaceous fern Onychiopsis tion. Cretaceous Research 29, 725e753. psilotoides from Bornholm, Denmark. Review of Palaeobotany and Palynology Haywood, A.M., Valdes, P.J., Markwick, P.J., 2004. Cretaceous (Wealden) climates: 66, 47e63. a modelling perspective. Cretaceous Research 25, 303e311. Friis, E.M., Pedersen, K.R., 1996. Eucommiitheca hirsuta, a new pollen organ with He, T., Pausas, J.G., Belcher, C.M., Schwilk, D.W., Lamont, B.B, 2012. Fire-adapted Eucommiidites pollen from the Early Cretaceous of Portugal. Grana 35, 104e112. traits of Pinus arose in the fiery Cretaceous. New Phytologist. doi:10.1111/j.1469- Friis, E.M., Skarby, A., 1981. Structurally preserved angiosperm flowers from the 8137.2012.04079.x. Upper Cretaceous of southern Sweden. Nature 291, 484e486. Herendeen, P.S., 1991a. Lauraceous wood from the mid-Cretaceous Potomac Group Friis, E.M., Skarby, A., 1982. Scandianthus gen. nov., angiosperm flowers of saxi- of eastern North America: Paraphyllanthoxylon marylandense sp. nov. Review of fragalean affinity from the upper Cretaceous of southern Sweden. Annals of Palaeobotany and Palynology 69, 277e290. Botany 50, 569e583. Herendeen, P.S., 1991b. Charcoalified angiosperm wood from the Cretaceous of Friis, E.M., Crane, P.R., Pedersen, K.R., 1986. Floral evidence for Cretaceous chlor- eastern North America and Europe. Review of Palaeobotany and Palynology 70, anthoid angiosperms. Nature 320, 163e164. 225e239. Friis, E.M., Crane, P.R., Pedersen, K.R., 1988. Reproductive structures of Cretaceous Herendeen, P.S., Skog, J.E., 1998. Gleichenla chaloneri e a new fossil fern from the Platanaceae. Biologiske Skrifter 31, 5e55. Lower Cretaceous (Albian) of England. International Journal of Plant Sciences Friis, E.M., Pedersen, K.R., Crane, P.R., 1992. Esgueiria gen. nov., fossil flowers with 159, 870e879. combretaceous features from Portugal. Biologiske Skrifter, Det Kongelige Herendeen, P.S., Crepet, W.L., Nixon, K.C., 1993. Chloranthus-like stamens from the Danske Videnskabernes Selskab 41, 1e45. Upper Cretaceous of New Jersey. American Journal of Botany 80, 865e871. Friis, E.M., Pedersen, K.R., Crane, P.R., 1994. Angiosperm floral structures from the Early Herendeen, P.S., Magallon-Puebla, S., Lupia, R., Crane, P.R., Kobylinska, J., 1999. Cretaceous of Portugal. Plant Systematics and Evolution Supplement 8, 31e49. A preliminary conspectus of the Allon flora from the Late Cretaceous (Late S.A.E. Brown et al. / Cretaceous Research 36 (2012) 162e190 189

Santonian) of the central Georgia, USA. Annals of the Missouri Botanical Garden Otto-Bliesner, B.L., Upchurch, G.R. Jr., 1997. Vegetation-induced warming of high- 86, 407e471. latitude regions during the Late Cretaceous period. Nature 385, 804e807. Herman, A.B., Spicer, R.A., 2010. Mid-Cretaceous floras and climate of the Russian Passalia, M.G., 2007. A mid-Cretaceous flora from the Kachaike Formation, Pata- high Arctic (Novosibirsk Islands, northern Yakutiya). Palaeogeography, Palae- gonia, Argentina. Cretaceous Research 28, 830e840. oclimatology, Palaeoecology 295, 409e422. Pausas, J.G., Keeley, J.E., 2009. A burning story: the role of fire in the history of life. Hermanová, Z., Kvacek, J., 2010. Late Cretaceous Palaeoaldrovanda, not seeds of Bioscience 59, 593e601. a carnivorous plant, but eggs of an insectJournal of the National Museum Pedersen, K.R., Friis, E.M., Crane, P.R., 1993. Pollen organs and seeds with Decusso- (Prague). Natural History Series 179, 105e118. sporites Brenner from the Lower Cretaceous Potomac Group sediments of Hickey, L.J., Doyle, J.A., 1977. Early Cretaceous fossil evidence for angiosperm eastern USA. Grana 32, 273e289. evolution. Botanical Review 43, 2e104. Pedersen, K.R., von Balthazar, M., Crane, P.R., Friis, E.M., 2007. Early Cretaceous floral Hollick, C.A., 1906. The Cretaceous flora of southern New York and New England. structures and in situ tricolpate-striate pollen: new early eudicots from United States Geological Survey Monographs 50, 1e219. Portugal. Grana 46, 176e196. Hollick, C.A., Jeffrey, E.C., 1909. Studies of Cretaceous coniferous remains from Pole, M.S., Douglas, J.G., 1999. Bennettitales, Cycadales and Ginkgoales from the mid Kreischerville, New York. Memoirs of the New York Botanical Garden 3, 1e137. Cretaceous of the Eromanga Basin, Queensland, Australia. Cretaceous Research Horrell, M.A., 1991. Phytogeography and paleoclimatic interpretation of the Maes- 20, 523e538. trichtian. Palaeogeography, Palaeoclimatology, Palaeoecology 86, 87e138. Pole, M., Philippe, M., 2010. Cretaceous plant fossils of Pitt Island, the Chatham Hu, S., Dilcher, D.L., Schneider, H., Jarzeny, D.M., 2006. Eusporangiate ferns from the Group, New Zealand. Alcheringa 29, 231e263. Dakota Formation, Minnesota, U.S.A. International Journal of Plant Sciences 167, Pyne, S.J., Andrews, P.L., Laven, R.D., 1996. Introduction to Wildland Fire. J. Wiley and 579e589. Sons, New York, NY, 769 pp. Hudspith, V., Scott, A.C., Collinson, M.E., Pronina, N., Beeley, T., 2012. Evaluating the Quan, C., Sun, C., Sun, Y., Sun, G., 2009. High resolution estimates of paleo-CO2 levels extent to which wildfire history can be interpreted from inertinite distribution through the Campanian (Late Cretaceous) based on Ginkgo cuticles. Cretaceous in coal pillars: an example from the late Permian, Kuznetsk Basin, Russia. Research 30, 424e428. International Journal of Coal Geology 89, 13e25. Royer, D.L., Miller, I.M., Peppe, D.J., Hickey, L.J., 2010. Leaf economic traits from Keeley, J.E., 2009. Fire intensity, fire severity and burn severity: a brief review and fossils support a weedy habit for early angiosperms. American Journal of Botany suggested usage. International Journal of Wildland Fire 18, 116e126. 97, 438e445. Keeley, J.E., Pausas, J.G., Rundel, P.W., Bond, W.J., Bradstock, R.A., 2011. Fire as an Sander, P.M., 1987. Taphonomy of the Lower Permian Geraldine Bonebed in Archer evolutionary pressure shaping plant traits. Trends in Plant Science 16, 406e411. County, Texas. Palaeogeography, Palaeoclimatology, Palaeoecology 61, 221e236. Knobloch, E., Mai, D.H., 1991. Evolution of middle and Upper Cretaceous floras in Saward, S.A., 1992. A global view of Cretaceous vegetation. Geological Society of central and western Europe. Geologisches Jahrbuch Reihe A 134, 257e270. America, Special Paper 267, 17e35. Kraal, P., Slomp, C.P., Forster, A., Kuypers, M.M.M., 2010. Phosphorous cycling from Schönenberger, J., Pedersen, K.R., Friis, E.M., 2001. Normapolles flowers of fagalean the margin to abyssal depths in the proto-Atlantic during anoxic event 2. affinity from the Late Cretaceous of Portugal. Plant Systematics and Evolution Palaeogeography, Palaeoclimatology, Palaeoecology 195, 42e54. 226, 205e230. Kring, D.A., 2007. The Chicxulub impact event and its environmental consequences Schulte, P., Alegret, L., Arenillas, I., Arz, J.A., Barton, P.J., Bown, P.R., Bralower, T.J., at the Cretaceous-Tertiary boundary. Palaeogeography, Palaeoclimatology, Christeson, G.L., Claeys, P., Cockell, C.S., Collins, G.S., Deutsch, A., Goldin, T.J., Palaeoecology 255, 4e21. Goto, K., Grajales-Nishimura, J.M., Grieve, R.A.F., Gulick, S.P.S., Johnson, K.R., Kump, L., 1988. Terrestrial feedback in atmospheric oxygen regulation by fire and Kiessling, W., Koeberl, C., Kring, D.A., MacLeod, K.G., Matsui, T., Melosh, J., phosphorous. Nature 335, 152e154. Montanari, A., Morgan, J.V., Neal, C.R., Nichols, D.J., Norris, R.D., Pierazzo, E., Kvacek, J., Eklund, H., 2003. A report on newly recovered reproductive structures Ravizza, G., Rebolledo-Vieyra, M., Reimold, W.U., Robin, E., Salge, T., Speijer, R.P., from the Cenomanian of Bohemia (central Europe). International Journal of Sweet, A.R., Urrutia-Fucugauchi, J., Vajda, V., Whalen, M.T., Willumsen, P.S., Plant Sciences 164, 1021e1039. 2010. The Chicxulub asteroid impact and mass extinction at the Cretaceous- Kvacek, J., Friis, E.M., 2010. Zlatkocarpus gen. nov., a new angiosperm reproductive Paleogene boundary. Science 327, 1214e1218. structure with monocolpate-reticulate pollen from the Late Cretaceous (Cen- Scott, A.C., 2000. The pre-Quaternary history of fire. Palaeogeography, Palae- omanian) of the Czech Republic. Grana 49, 115e127. oclimatology, Palaeoecology 164, 281e329. Leng, Q., Schönenberger, J., Friis, E.M., 2005. Late Cretaceous follicular fruits from Scott, A.C., 2010. Charcoal recognition, taphonomy and uses in palaeoenvironmental southern Sweden with systematic affinities to early diverging dicots. Botanical analysis. Palaeogeography, Palaeoclimatology, Palaeoecology 291, 11e39. Journal of the Linnean Society 148, 377e407. Scott, A.C., Collinson, M.E., 1978. Organic sedimentary particles: results from SEM of Lenton, T.M., Watson, A.J., 2000. Redfield revisited: 2. What regulates the oxygen fragmentary plant material. In: Whalley, W.B. (Ed.), SEM in the Study of Sedi- content of the atmosphere? Global Biogeochemical Cycles 14, 249e268. ments. Geo Abstracts, Norwich, pp. 137e167. Lidgard, S., Crane, P.R.,1990. Angiosperm diversification and Cretaceous floristic trends Scott, A.C., Glasspool, I.J., 2007. Observations and experiments on the origin and e a comparison of palynofloras and leaf macrofloras. Paleobiology 16, 77e93. formation of inertinite group macerals. International Journal of Coal Geology Lupia, R., 2011. Late Santonian megaspore floras from the Gulf Coastal Plain 70, 53e66. (Georgia, USA). Journal of Paleontology 85, 1e21. Scott, A.C., Jones, T.P., 1994. The nature and influence of fires in Carboniferous Lupia, R., Lidgard, S., Crane, P.R., 1999. Comparing palynological abundance and ecosystems. Palaeogeography, Palaeoclimatology, Palaeoecology 106, 91e112. diversity: implications for biotic replacement during the Cretaceous angio- Scott, A.C., Stea, R., 2002. Fires sweep across the mid-Cretaceous landscape of Nova sperm radiation. Paleobiology 25, 305e340. Scotia. Geoscientist 12, 4e6. Martill, D.M., Loveridge, R.F., Mohr, B.A.R., Simmonds, E., 2012. A wildfire origin for Scott, A.C., Cripps, J., Nichols, G., Collinson, M.E., 2000. The taphonomy of charcoal terrestrial organic debris in the Cretaceous Santana Formation Fossil Lagerstätte following a recent heathland fire and some implications for the interpretation of (Araripe Basin) of north-east Brazil. Cretaceous Research 34, 135e141. fossil charcoal deposits. Palaeogeography, Palaeoclimatology, Palaeoecology 164, Mendes, M.M., Pais, J., Pedersen, K.R., Friis, M., 2010. Erdtmanitheca portucalensis, 1e31. a new pollen organ from the Early Cretaceous (AptianeAlbian) of Portugal with Sellwood, B.W., Valdes, P.J., 2006. Mesozoic climates: general circulation models Eucommiidites-type pollen. Grana 49, 26e36. and the rock record. Sedimentary Geology 190, 269e287. Miller, C.N., 1999. Implications of fossil conifers for the phylogenetic relationships of Sender, L.M., Diez, J.B., Ferrer, J., Pons, D., Rubio, C., 2005. Preliminary data on a new living families. The Botanical Review 65, 239e277. Albian flora from the Valle del Río Martín, Teruel, Spain. Cretaceous Research Moody, J.A., Martin, D.A., 2001. Initial hydrologic and geomorphic response 26, 898e905. following a wildfire in the Colorado Front Range. Earth Surface Processes and Sewall, J.O., Van De Wal, R.S.W., van der Zwan, K., van Oosterhout, C., Dijkstra, H.A., Landforms 26, 1049e1070. Scotese, C.R., 2007. Climate model boundary conditions for four Cretaceous time Moody, J.A., Martin, D.A., 2009. Forest fire effects on geomorphic processes. In: slices. Climate of the Past 3, 647e657. Cerdá, A., Robichaud, P. (Eds.), Fire Effects on Soils and Restoration Strategies. Shakesby, R.A., Doerr, S.H., 2006. Wildfire as a hydrological and geomorphological Science Publishers, Inc, Enfield, NH, pp. 41e79. agent. Earth-Science Reviews 74, 269e307. Moody, J.A., Martin, D.A., Cannon, S.H., 2008. Post-wildfire erosion response in two Simoneit, B.R.T., 2002. Biomass burning e a review of organic tracers for smoke geologic terrains in the western USA. Geomorphology 95, 103e118. from incomplete combustion. Applied Geochemistry 17, 129e162. Najarro, M., Peñalver, E., Pérez-de-la Fuente, P., Ortega-Blanco, J., Menor-Salván, C., Sims, H.J., Herendeen, P.S., Crane, P.R., 1998. New genus of fossil Fagaceae from the Barróne, E., Soriano, C., Rosales, I., López del Valle, R., Velasco, F., Torrnos, F., Santonian (Late Cretaceous) of central Georgia, U.S.A. International Journal of Daviero-Gomez, V., Gomez, B., Delclòs, X., 2010. Review of the El Soplao amber Plant Sciences 159, 391e404. outcrop, Early Cretaceous of Cantabria, Spain. Acta Geologica Sinica 84, 959e976. Sims, H.J., Herendeen, P.S., Lupia, R., Christopher, R.A., Crane, P.R., 1999. Fossil Nicholas, C.J., Henwood, A.A., Simpson, M., 1993. A new discovery of early Cretaceous flowers with Normapolles pollen from the Upper Cretaceous of southeastern (Wealden) amber from the Isle of Wight. Geological Magazine 130, 847e850. North America. Review of Palaeobotany and Palynology 106, 131e151. Niklas, K.J., Tiffany, B.H., Knoll, A.H., 1985. Patterns in vascular plant diversification: an Spicer, R.A., 2003. Changing climate and biota. In: Skelton, P. (Ed.), The Cretaceous analysis at the species level. In: Valentine, J.W. (Ed.), Phanerozoic Diversity Patterns: World. Cambridge University Press, Cambridge, pp. 85e163. Profiles in Macroevolution. Princeton University Press, Princeton, NJ, pp. 97e128. Spicer, R.A., Corfield, R.M., 1992. A review of terrestrial and marine climates in the Nixon, K.C., Crepet, W.L., 1993. Late Cretaceous fossil flowers of Ericalean affinity. Cretaceous with implications for modeling the ‘Greenhouse Earth’. Geological American Journal of Botany 80, 616e623. Magazine 129, 169e180. Nyman, P., Sheridan, G., Lane, P.N.J., 2010. Synergistic effects of water repellency and Spicer, R.A., Herman, A.B., 2001. The AlbianeCenomanian flora of the Kukpowruk macropore flow on the hydraulic conductivity of a burned forest soil, south-east River, western North Slope, Alaska: stratigraphy, palaeofloristics, and plant Australia. Hydrological Processes 24, 2871e2887. communities. Cretaceous Research 22, 1e40. 190 S.A.E. Brown et al. / Cretaceous Research 36 (2012) 162e190

Spicer, R.A., Ahlberg, A., Herman, A.B., Hofmann, C.C., Raikevich, M., Valdes, P.J., Váchová, V., Kvacek, J., 2009. Palaeoclimate analysis of the flora of the Klikov Marwick, P.J., 2008. The Late Cretaceous continental interior of Siberia: a chal- Formation, Upper Cretaceous, Czech Republic. Bulletin of Geosciences 84, lenge for climate models. Earth and Planetary Science Letters 267, 228e235. 257e268. Srinivasan, V., 1992. Two new species of the conifer Glenrosa from the Lower Creta- von Balthazar, M., Pedersen, K.R., Friis, E.M., 2005. Teixeiria lusitanica, a new fossil ceous of North America. Review of Palaeobotany and Palynology 72, 245e255. flower from the Early Cretaceous of Portugal with affinities to Ranunculales. Sweetman, S.C., Insole, A.N., 2010. The plant debris beds on the Early Cretaceous Plant Systematics and Evolution 255, 55e75. (Barremian) Wessex Formation of the Isle of Wight, southern England: their von Balthazar, M., Pedersen, K.R., Crane, P.R., Stampanoni, M., Friis, E.M., 2007. genesis and palaeontological significance. Palaeogeography, Palaeoclimatology, Potomacanthus lobatus gen. et. sp. nov., a new flower of probable Lauraceae Palaeoecology 292, 409e424. from the Early Cretaceous (Early to Middle Albian) of eastern North America. Takahashi, M., Crane, P.R., Ando, H., 1999a. Fossil flowers and associated plant fossils American Journal of Botany 94, 2041e2053. from the Kamikitaba locality (Ashizawa Formation, Futaba Group, Lower Con- Wang, X., 2004. Plant cytoplasm preserved by lightning. Tissue and Cell 36, 351e360. iacian, Upper Cretaceous) of Northeast Japan. Journal of Plant Research 112, Watson, J., Alvin, K.L., 1996. An English Wealden floral list, with comments on 187e206. possible environmental indicators. Cretaceous Research 17, 5e26. Takahashi, M., Crane, P.R., Ando, H., 1999b. Esguieria futabensis sp. nov., a new Whitlock, C., Higuera, P.E., McWethy, D.B., Briles, C.E., 2010. Paleoecological angiosperm flower from the Upper Cretaceous (Lower Coniacian) of north- perspectives on fire ecology: revisiting the fire-regime concept. The Open eastern Honshu, Japan. Palaeontological Research 3, 81e87. Ecology Journal 3, 6e23. Takahashi, M., Friis, E.M., Herendeen, P.S., Crane, P.R., 2008. Fossil flowers of Fagales Wildman, R.A., Hickey, L.J., Dickinson, M.B., Berner, R.A., Robinson, J.M., Dietrich, M., from the Kamikitaba locality (Early Coniacian; Late Cretaceous) of northeastern Essenhigh, R.H., Wildman, C.B., 2004. Burning of forest materials under late Japan. International Journal of Plant Sciences 169, 899e907. Paleozoic high atmospheric oxygen levels. Geology 32, 457e460. Taylor, D.W., Hickey, L.J., 1996. Evidence for and implications of an herbaceous origin Wing, S.L., Boucher, L.D., 1998. Ecological aspects of the Cretaceous flowering plant for angiosperms. In: Taylor, D.W., Hickey, L.J. (Eds.), Flowering Plant Origin, radiation. Annual Review Earth Planetary Science 26, 379e421. Evolution and Phylogeny. Chapman and Hall, New York, NY, pp. 232e266. Wolbach, W.S., Gilmour, I., Anders, E., 1990. Major wildfires at the K-T boundary. In: Tiffney, B.H., 1977. Dicotyledonous angiosperm flower from the Upper Cretaceous of Sharpton, V.L., Ward, P.D. (Eds.), Global catastrophies in Earth History, 247. Martha’s Vineyard, Massachusetts. Nature 265, 136e137. Geological Society of America, Special Paper, pp. 391e400. Troost, G., 1821. Description of a variety of amber and of a fossil substance supposed Wolfe, J.A., Upchurch, G.R. Jr., 1987. North American nonmarine climates and to be the nest of an insect discovered at Cape Sable, Magothy River, Ann- vegetation during the Late Cretaceous. Palaeogeography, Palaeoclimatology, Arundel Country, Maryland. American Journal of Science and Arts 3, 8e15. Palaeoecology 61, 33e77. Ulicný, D., Kvacek, J., Svobodová, M., Spi cáková, L., 1997. High-frequency sea-level Wuyep, E.O., Obaje, N.G., 2010. Petrographic evaluation of the ranks and techno- fluctuations and plant habitats in Cenomanian fluvial to estuarine succession: logical applications of some coal deposits in the Anambra Basin and Middle Pecínov Quarry, Bohemia. Palaeogeography, Palaeoclimatology, Palaeoecology Benue Trough of Nigeria. Geological Society of America. Abstracts with 136, 165e197. Programs, Houston, TX.